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Review of Janenschia Wild, with the description of a new sauropod from the Tendaguru beds of Tanzania and a discussion on the systematic value of procoelous caudal vertebrae in the sauropoda
Abstract
Since the holotype of Janenschia robusta (E. FRAAS) is based on an incomplete right hind limb, only specimens with comparable parts of the skeleton can be properly referred to that species. This is the case with the material from Tendaguru site P (JANENSCH 1925, 1929a), which consists of fore and hind limb skeletons from two individuals. The remaining sauropod skeletal material, i. e. the isolated anterior dorsal vertebrae, an articulated tail, and an anterior caudal vertebra, collected in the surroundings of Tendaguru Hill (JANENSCH 1929a) can not be properly referred to Janenschia due to the incompleteness of the type material of Janenschia robusta (E. FRAAS). Janenschia robusta (E. FRAAS) is redescribed and diagnosed, on the basis of the holotype and referred specimens from Tendaguru site P. Morphological comparisons suggest closer relationships between Janenschia and Camarasaurus. A new genus and species, Tendaguria tanzaniensis gen. et sp. nov. is proposed for two anterior dorsal vertebrae collected at Nambango (JANENSCH 1929a). Tendaguria gen. nov. is mainly diagnosed by extremely reduced neural spines which are not higher than the neural arch. They are connected with the postzygapophysial laminae and the epipophyses building a common dorsal plane. Comparison of these anterior dorsal vertebrae with those of different sauropod genera suggests that the vertebrae of Tendaguria gen. nov. represent the more derived type. The neural arch pattern in Tendaguria tanzaniensis gen. et sp. nov. points to a strong modification in the position and function of the episomatic musculature whose origin and insertion are both on these vertebrae. An articulated tail consisting of 30 caudal vertebrae collected at site G near Tendaguru Hill (JANENSCH 1929a), formerly referred to Janenschia and considered to have titanosaurid affinities, is described and compared to other titanosaurs. This caudal vertebrae series exhibits decreasing procoely posteriorly. The first 10 vertebrae are procoelous, the following 3 caudals are nearly amphiplatyan, and the last 17 are slightly amphicoelous. The procoelous condition in anterior caudal vertebrae is also known in Jurassic sauropods from China (e. g. Bellusaurus, Mamenchisaurus). This vertebral type was developed independently in different sauropod lineages. In the Late Cretaceous family Titanosauridae, all caudal vertebrae are characterized by procoely. It is concluded that sauropod families can not be distinguished by their procoelous anterior caudal vertebrae.
... The wrinkles are more prominent on the lingual surface of the Jaisalmer specimen whereas, wrinkles are more prominent on the labial surface of the Turiasaurus and Losillasaurus. The SI value is lower than the average SI values of Mierasaurus, Moabo saurus, Narindasaurus and Turiasaurus and almost equal to Zby, but well within the range of Losillasau rus and the Portuguese heart-shaped teeth (Bonaparte et al. 2000;Royo-Torres et al. 2006;Mannion 2010;Mateus et al. 2014;Britt et al. 2017;Royo-Torres et al. 2017;Mannion et al. 2019;Royo-Torres et al. 2021). The Jaisalmer crown has a concave lingual sur-face unlike Cardiodon (Upchurch & Martin 2003). ...
... Turiasauria clade includes the Middle Jurassic Narin dasaurus from Madagascar, the Late Jurassic Tend aguria from Africa, the Late Jurassic Losillasaurus, Turiasaurus, Zby and Amanzia from Europe, and the Early Cretaceous Mierasaurus and Moabosaurus from North America (Bonaparte et al. 2000;Royo-Torres et al. 2006;Mannion 2010;Mateus et al. 2014;Britt et al. 2017;Royo-Torres et al. 2017;Mannion et al. 2019;Schwarz et al. 2020;Royo-Torres et al. 2021). Turiasaur remains are found from the Middle Jurassic to the Cretaceous. ...
... ported exclusively in Turiasauria clade(Royo-Torres et al. 2021). Turiasauria is a stem-based clade, formed byRoyo-Torres et al. (2006) for all taxa comparatively more related to Turiasaurus than Saltasaurus.As per the phylogenetic analysis carried out byRoyo- Torres et al. (2021), this clade includes Turiasaurus,Mierasaurus, Losillasaurus, Narindasaurus, Zby, Moabosaurus, Tendaguria, and probably Amanzia (Royo-Torres et al. 2006;Mannion 2010;Bonaparte et al. 2000;Mateus et al. 2014;Britt et al. 2017;Royo-Torres et al. 2017;Mannion et al. 2019;Schwarz et al. 2020;Royo-Torres et al. 2021). In heart-shaped morphology, the distal and the mesial edges of the broad crown converge towards the apex in the apical half with maximum width at the base of the apex which is almost halfway from apex to base. ...
This study presents the dental record of a non-neosauropod eusauropod from the Middle Jurassic rocks of the Jaisalmer Basin, India. An isolated, fragmentary heart-shaped crown of a turiasaur has been recovered from the intraformational conglomerate of the Badabag Member (Bathonian) of the Jaisalmer Formation. The heart-shaped, asymmetrical crowns with a labio-lingually compressed apex and apicobasal shallow grooves in the root are exclusive to the Turiasauria clade. This is the first record of a turiasaur from India as well as from Asia. This study extends the geographic distribution of the Turiasauria clade and provides new data about the diversity in the Jurassic sauropod fauna of India. The presence of heart-shaped teeth, from both Laurasia and Gondwanaland, shows a much wider geographic and stratigraphic distribution of the Turiasauria clade. There was a possibility of faunal exchange between the two big landmasses at least during the Jurassic. This study is one more step in bringing out the diversity in the vertebrate assemblage of the Jaisalmer Basin during the Middle Jurassic.
... It is likely that on the lateral face of the proximal end, there is a triangular scar for the articulation of the fibula. There are a sign of a second cnemial crest sensu Bonaparte et al. (2000) and Mannion et al. (2013). Here the feeble second cnemial crest is observed on the anterior side of fibular articular condylar ridge (Fig.42) like Lusotitan atalaiensis Mannion et al. (2013) and Gspsaurus from Indo-Pakistan (present research), and unlike Oceanotitan dantasi Mocho et al. (2019), Pakisaurus and Isisaurus from IndoPakistan (present research). ...
... On the anterior part of lateral face of the proximal end, there is a triangular scar for the articulation of the fibula. There are a sign of a second cnemial crest (Fig.4) sensu Bonaparte et al. (2000) andMannion et al. (2013). Here the well developed second cnemial crest (Fig.4) is observed on the anterior side of fibular articular condylar ridge like Lusotitan atalaiensis Mannion et al. (2013) and Saraikimasoom vitakri from Indo-Pakistan (present research), and unlike Oceanotitan dantasi Mocho et al. (2019), Pakisaurus balochistani and Isisaurus colberti from Indo-Pakistan (present research). ...
... Bellusaurus (Dong, 1990), Chuanjiesaurus (Sekiya, 2011), HMN MB.R.2091.1-30 ( Bonaparte et al., 2000;Mannion et al. 2013), andLosillasaurus (Casanovas, Santafe & Sanz, 2001)]. Similarly, the extension of procoely into middle-posterior caudal vertebrae has been considered a feature of lithostrotian titanosaurs (Jacobs et al., 1993;Upchurch, 1995Upchurch, , 1998Wilson, 2002). ...
... The proximal surface bears a boss near the lateral edge and above the fibular articulation. There are no signs of a second cnemial crest sensu Bonaparte et al. (2000) and Mannion et al. (2013). The diaphysis is also transversely compressed, probably due to taphonomic effects, with an oval outline in cross-section (the maximum diameter is anteroposteriorly oriented). ...
... Nevertheless, this taxon was recently recovered as a diplodocine diplodocid by Tschopp et al. (2015). Janenschia is another possible somphospondylan taxon with a relatively complex taxonomic history (see Bonaparte et al., 2000;Mannion et al., 2013). This taxon was considered to be a basally branching titanosaur (Upchurch, 1995;Upchurch et al., 2004) in disagreement with Bonaparte et al. (2000), who considered it to be a form closely related to Camarasaurus. ...
... Janenschia is another possible somphospondylan taxon with a relatively complex taxonomic history (see Bonaparte et al., 2000;Mannion et al., 2013). This taxon was considered to be a basally branching titanosaur (Upchurch, 1995;Upchurch et al., 2004) in disagreement with Bonaparte et al. (2000), who considered it to be a form closely related to Camarasaurus. More recent works support Janenschia as a non-titanosauriform macronarian (Carballido et al., 2011;Mannion et al., 2013Mannion et al., , 2017Carballido and Sander, 2014;Upchurch et al., 2015; this work; and also in agreement with D'Emic, 2012). ...
The Upper Jurassic of Portugal is relatively rich in sauropod remains. We describe a new sauropod specimen, which includes a partial tail, pectoral and pelvic girdle elements, and hind limb bones, from Praia de Valmitão (Praia da Amoreira-Porto Novo Formation, upper Kimmeridgian–lowermost Tithonian). This specimen constitutes the holotype of Oceanotitan dantasi, gen. et sp. nov., which shows a unique combination of characters, including the presence of anterior caudal vertebrae with a medial accessory articulation on the prezygapophysis; a circular, rough tuberosity on the medial surface of the scapula; an elliptical concavity on the ventral face of the scapula; an ischium that is shorter than the pubis; and a robust fourth trochanter located at the midline of the posterior face of the femur. Multiple phylogenetic analyses recover Oceanotitan dantasi within Titanosauriformes, with one resolving it at the base of Somphospondyli. This taxon shares several apomorphies with some Cretaceous somphospondylans and turiasaurs, such as the transverse furrow on the chevron articulations (shared with Tangvayosaurus and Phuwiangosaurus) and the ischium being shorter than the pubis (shared with Mierasaurus and somphospondylans). Oceanotitan might represent the oldest known somphospondylan, and its establishment increases the known diversity of the Late Jurassic–earliest Early Cretaceous sauropod fauna in the Iberian Peninsula, which also consists of turiasaurs, diplodocids and macronarians (non-camarasaurid, non-titanosauriform macronarians; camarasaurids; and brachiosaurids). This high diversity in sauropods suggests that this region might have played an important role during the Late Jurassic in the dispersal and diversification of several sauropod lineages between North America, Africa, and Europe, especially macronarians.
... The convexity of the posterior articulation is more pronounced than in MNHNUL/P.R27, SHN 180 and MMPM.P/73, which also suggest a more anterior position in the tail. The presence of a more pronounced posterior convexity in the anteriormost caudal vertebrae also occurs in other sauropods with this type of articulation, including non-neosauropods eusauropods (Bonaparte et al., 2000;Casanovas et al., 2001;Fig. 3. Anterior caudal vertebra of Praia da Corva (Torres Vedras, SHN 530) in anterior (a), left (b), ventral (c), posterior (d), right (e), and dorsal (f) views. ...
... Description: An anterior caudal vertebra was recovered from the cliffs of Baleal (Peniche). Taking into account the presence of a deep caudal rib extending to the lateral surface of the neural arch, and comparing with other well-known caudal series (e.g., Osborn and Mook, 1921;Bonaparte et al., 2000), this caudal vertebra probably corresponds to an anterior one, between the fourth and eighth caudal vertebrae. SHN 180 occupies a more posterior position than SHN 530, MNHNUL/P.R27 and MMPM.P/73. ...
... MB.R.2091.1-30 is a caudal series referred by Janensch (1929) to Gigantosaurus robustus (now Janenschia robustus) from the Tendaguru Fm. (Upper Jurassic), which also presents strong-toslight procoelous caudal vertebrae (Bonaparte et al., 2000). The higher neural spines and the anterolaterally directed caudal ribs differentiate MB.R.2091.1-30 ...
The Upper Jurassic of the Lusitanian Basin (Portugal) is particularly rich in sauropod fossil remains, with four established taxa: Dinheirosaurus, Lusotitan, Lourinhasaurus and Zby. The presence of sauropod caudal procoelous vertebrae is reported for the first time in the Upper Jurassic of Portugal, with specimens described from the localities of Baleal, Paimogo, Praia da Areia Branca, Porto das Barcas, and Praia da Corva. The presence of slightly procoelous centra and fan-shaped caudal ribs with smooth prezygapophyseal centrodiapophyseal fossa in the more anterior caudal vertebrae allows for the assignment of these specimens to an indeterminate eusauropod, probably belonging to a non-neosauropod eusauropod form. The absence of several features in the Portuguese specimens that are common in diplodocids, mamenchisaurids and titanosaurs, prevents the establishment of sound relationships with these clades. The described specimens are almost identical to the anterior caudal vertebrae of the Iberian turiasaur Losillasaurus. During the Iberian Late Jurassic, Turiasauria is the only Iberian group of sauropods which shares this type of morphology with the Baleal, Paimogo, Praia da Areia Branca, Porto das Barcas and Praia da Corva specimens. These specimens represent one of the four anterior caudal vertebral morphotypes recorded in the Upper Jurassic of the Lusitanian Basin and briefly described herein.
... Turiasuria is a clade of large-bodied, eusauropod dinosaurs erected by Royo-Torres et al. (2006) [1], hitherto known from the Middle Jurassic to Early Cretaceous of Europe, North America, Africa, and India [1][2][3][4][5][6][7][8]. Turiasaurs have distinct heart-shaped teeth, which are exclusive to their clade and easily recognizable (character 402 [6]). ...
... Temporal distribution of the different recognized members of the Turiasauria, based on ages given by[1][2][3][4][5][6][7][8]. ...
Turiasauria is a clade of basal sauropod dinosaurs hitherto only known from the Middle Jurassic (Bathonian) to the Lower Cretaceous (Valanginian). A new find of a shed tooth crown from the Lower Jurassic (Pliensbachian), Halse Formation of Bornholm, Denmark, is spoon-like, asymmetrical, and heart-shaped, which identifies the tooth as turiasaurian, pushing the origin of the Turiasauria some 17 My back into the Lower Jurassic. This suggests a North Pangean/Laurasian origin of the turiasaurian clade, which then, during the Middle to Late Jurassic, dispersed through Europe, India, and Africa, with their latest representatives found in the Early Cretaceous of England and North America. Furthermore, this is the first record of a sauropod from the Pliensbachian in Europe.
... A small ridge posterior to the cnemial crest is present at the top of this groove. This crest is interpreted as the second cnemial crest described by Bonaparte et al. (2000) for the tibia (SMNS 12144) of Janenschia. In the San Lorenzo specimen, this crest is markedly pointed and parallel to the cnemial crest. ...
... We have two possible groups of turiasaurs: one for Narindasaurus as a sistertaxon of the European turiasaurs during the Jurassic, with Amanzia, Losillasaurus, Turiasaurus and Zby, and another clade for the Late Jurassic and Early Cretaceous turiasaurs from Africa and North America. The latter clade includes Tendaguria with Moabosaurus and Mierasaurus as a second family of turiasaurs, which might imply the recovery of the Tendaguriidae clade established by Bonaparte et al. (2000). However, we prefer to wait until the relationships are tested with new data and further analyses, as suggested by Mannion et al. (2019). ...
Turiasauria is a non-neosauropod eusauropod clade of dinosaurs known since 2006, when the description of Turiasaurus was published. This group, including Losillasaurus, was originally thought to have been restricted to the Late Jurassic of Spain. However, over the last decade, our knowledge of this group has improved with the discovery of new taxa such as Zby from the Portuguese Late Jurassic, Tendaguria from the Tanzanian Late Jurassic and Mierasaurus and Moabosaurus from the Early Cretaceous of the USA. Here, we describe a new specimen of Losillasaurus from Spain, which allows us to better understand the character variation in the cranial and postcranial skeleton. The review of some sauropod fauna of Madagascar, and inclusion of some specimens of Turiasauria, suggest that this clade might have arisen in the Middle Jurassic. According to our phylogenetic results, a specimen found in the early 19th century in Madagascar is shown to be the oldest and only member of Turiasauria represented in the Middle Jurassic thus far. This is named Narindasaurus thevenini gen. & sp. nov.. Turiasauria is thus known from the Middle Jurassic in Pangaea, diversified in the Late Jurassic in Gondwana and Laurasia, and dispersed during the Early Cretaceous to North America.
... A claviform neural spine on the anteriormost caudal vertebrae is likewise present in Chuanjiesaurus (LCD9701-I), the holotype, but not the referred specimen, of Mamenchisaurus hochuanensis (CCG V 20401; ZDM 0126) and Wamweracaudia keranjei (HMN MB.R.2091.1-30; Janensch 1929b; Bonaparte et al. 2000;Mannion et al. 2019), although in the latter case the SPRLs and SPOLs are more fully developed and make the margins of the spine more distinct. By contrast, Mamenchisaurus youngi (Ouyang & Ye 2002, fig. ...
... It does not extend far laterally, but obscures the weak anterior crest of the fibula in anterior view. The proximal lateral edge of the tibia forms a weak lip that overhangs the internal fossa for the fibula; no pinched projection -the 'second cnemial process' (Bonaparte et al. 2000) -is present along this edge. The second cnemial process is also absent in Bellusaurus (IVPP V17768), Euhelopus, Jobaria, Janenschia and various neosauropods (Mannion et al. 2013(Mannion et al. , 2017(Mannion et al. , 2019. ...
Fossil-rich deposits from the Middle and Late Jurassic of China have yielded a diverse array of sauropod dinosaurs, including numerous species referred to Mamenchisaurus and Omeisaurus. Despite an abundance of fossils and a proliferation of taxa, the anatomy of Middle–Late Jurassic Chinese sauropods remains poorly documented. Here, we comprehensively redescribe and illustrate Klamelisaurus gobiensis from the Middle–Late Jurassic Shishugou Formation of northwest China. Phylogenetic analyses conducted under parsimony and time-calibrated Bayesian optimality criteria consistently recover Klamelisaurus as a member of a predominantly Chinese radiation of exceptionally long-necked eusauropods that includes Mamenchisaurus spp., Chuanjiesaurus, Qijianglong and Wamweracaudia. In most analyses, this lineage also includes Euhelopus, reviving a ‘traditional’ Euhelopodidae and calling into question the macronarian affinities of Euhelopus. Klamelisaurus shares several features with Euhelopus that are unique to a subset of East Asian taxa or rare among sauropods, including a convex ventral margin of the prezygodiapophyseal lamina in middle–posterior cervical vertebrae, a ventrally bifurcated postzygodiapophyseal lamina in posterior cervical vertebrae, and development of a rugose projection extending anteriorly from the epipophysis into the spinodiapophyseal fossa in most cervical vertebrae. Anatomical comparisons of the cervical vertebrae of Klamelisaurus to several other sauropodomorphs and insights from myological studies of extant archosaurs strongly suggest that this latter structure, often considered part of an epipophyseal-prezygapophyseal lamina, is an epaxial muscle scar that is distinct from pneumatic structures of the lateral surface of the neural spine. The phylogenetic and comparative anatomical data presented here provide a foundation for future revision of the taxonomy and systematics of sauropods from the Junggar and Sichuan basins.
... Importantly, other major sauropod groups (e.g., rebbachisaurids, dicraeosaurids, euhelopodids, brachiosaurids) persisted alongside early titanosaurians from the Early Cretaceous until all non-titanosaurian sauropods finally succumbed to extinction by the early Late Cretaceous with their fossil remains recovered from North and South America, northern Africa, Europe, and Asia [9,20,22,[24][25][26][27][28][29][30]. With many different sauropod clades present globally during the Early Cretaceous, some of which clearly represent groups with a Jurassic origin, the origins of titanosaurians remain elusive among the diverse assemblage of sauropod clades known from around the world [17,29,[31][32][33]. ...
... The cnemial crest is directed anteriorly with a slight lateral curve (Fig 25E), as is exhibited in other titanosauriforms like the euhelopodid P. sirindhornae, the titanosauriform T. sanzi, and the titanosaurians R. krausei and M. dixeyi [33]. The cnemial crest is concave along its lateral surface to accommodate the proximal fibula and there is no second cnemial crest (i.e., an additional articulating ridge) as in some somphospondylian titanosauriforms [31,33]. The anterior margin of the cnemial crest is rugose along its medial edge. ...
The African terrestrial fossil record has been limited in its contribution to our understanding of both regional and global Cretaceous paleobiogeography, an interval of significant geologic and macroevolutionary change. A common component in Cretaceous African faunas, titanosaurian sauropods diversified into one of the most specious groups of dinosaurs worldwide. Here we describe the new titanosaurian Mnyamawamtuka moyowamkia gen. et sp. nov. from the Mtuka Member of the Galula Formation in southwest Tanzania. The new specimen preserves teeth, elements from all regions of the postcranial axial skeleton, parts of both appendicular girdles, and portions of both limbs including a complete metatarsus. Unique traits of M. moyowamkia include the lack of an interpostzygapophyseal lamina in posterior dorsal vertebrae, pronounced posterolateral expansion of middle caudal centra, and an unusually small sternal plate. Phylogenetic analyses consistently place M. moyowamkia as either a close relative to lithostrotian titanosaurians (e.g., parsimony, uncalibrated Bayesian analyses) or as a lithostrotian and sister taxon to Malawisaurus dixeyi from the nearby Aptian? Dinosaur Beds of Malawi (e.g., tip-dating Bayesian analyses). M. moyowamkia shares a few features with M. dixeyi, including semi-spatulate teeth and a median lamina between the neural canal and interpostzygapophyseal lamina in anterior dorsal vertebrae. Both comparative morphology and phylogenetic analyses support Mnyamawamtuka as a distinct and distant relative to Rukwatitan bisepultus and Shingopana songwensis from the younger Namba Member of the Galula Formation with these results largely congruent with newly constrained ages for the Mtuka Member (Aptian–Cenomanian) and Namba Member (Campanian). Coupled with recent discoveries from the Dahkla Oasis, Egypt (e.g., Mansourasaurus shahinae) and other parts of continental Afro-Arabia, the Tanzania titanosaurians refine perspectives on the development of African terrestrial faunas throughout the Cretaceous—a critical step in understanding non-marine paleobiogeographic patterns of Africa that have remained elusive until the past few years.
... The rhomboidal cross section makes the tibia of Tastavinsaurus closely resemble that of Brachiosaurus (Janensch, 1961). The groove on the cnemial crest has only been reported in Janenschia (Bonaparte et al., 2000), Phuwiangosaurus also has a ventral depression in its cnemial crest (Martin et al., 1999) that might be a similar structure. The twisted tibia is shared with other members of the Titanosauriformes possibly representing a synapomorphy of the clade. ...
... Such a high value has otherwise only been found in an isolated foot from the Early Cretaceous of Siberia belonging to an unidentified titanosauriform (Averianov et al., 2002). Metatarsal I is shorter than metatarsal V, as in other titanosauriforms (Gallup, 1989), which distinguishes it from basal macronarians such as Camarasaurus (Ostrom and McIntosh, 1966) or Janenschia (Bonaparte et al., 2000) in which metatarsal I is longer than metatarsal V. ...
The new sauropod dinosaur Tastavinsaurus sanzi, gen. et sp. nov., from the early Aptian of Spain is described. The holotype is a partially articulated skeleton of an adult individual recovered from the Arsis-1 site in Peñarroya de Tastavins (Teruel) at the base of the marine Xert Formation. It is one of the most complete and best-preserved sauropod dinosaur skeletons from the European Early Cretaceous. The fossil remains comprise the three caudalmost thoracic vertebrae, part of a fourth, nine thoracic rib fragments, sacrum, 25 caudal vertebrae, 21 chevrons, both ilia, pubes, ischia and femora, right tibia, right fibula, six metatarsals, and seven pedal phalanges (including four unguals). The new taxon is defined by 19 autapomorphies. In our cladistic analysis, Tastavinsaurus is the sister-taxon of the North American Venenosaurus within Titanosauriformes, which includes the Brachiosauridae, Somphospondyli, and Titanosauria. The new taxon provides new information about the diversity of non-brachiosaurid titanosauriforms during the Early Cretaceous in Europe and paleobiogeographic relationships between Europe and North America.
... 'Allosaurus tendagurensis ' Janensch, 1925 'Ceratosaurus roechiingi ' Janensch, 1925 'Labrosaurus stechowi ' Janensch, 1920 'Megalosaum ingens ' Janensch, 1920 Theropoda indet. Barosaurus africanus (Fraas, 1908) Barosaurus gracilis Janensch, 1961 Dicraeosaurus hansemanni Janensch, 1914c Dicraeosaurus sattleri Janensch, 1914c Brachiosaurus brancai Janensch, 1914c Janenschia robusta (Fraas, 1908) Tendaguria tanzaniensis Bonaparte et al., 2000 Sauropoda indet. ...
... The sauropods include Brachiosaurus brancai, Janenschia robusta, Dicraeosaurus sattleri and Barosaurus afvicanus (Janensch 1961). Fossiliferous deposits exposed at Nambango (approximately 15 km southeast of Tendaguru Hill) that probably belong to the Upper Saurian Bed yielded two anterior dorsal vertebrae of a sauropod assigned to Tendaguria tanzaniensis (Bonaparte et al. 2000). The ornithischians are represented by Kentrosaurus aethiopicus, the theropods by Elaphrosaurus bambergi, 'Megalosaurus ingens', 'Labrosaurus stechowi' and 'Ceratosaurus roechlingi' (Janensch 1925b, see also Table 2). ...
The Late Jurassic to Early Cretaceous Tendaguru Beds (Tanzania, East Africa) have been well known for nearly a century for their diverse dinosaur assemblages. Here, we present sedimentological and palaeontological data collected by the German-Tanzanian Tendaguru Expedition 2000 in an attempt to reconstruct the palaeo-ecosystems of the Tendaguru Beds at their type locality. Our reconstructions are based on sedimentological data and on a palaeoecological analysis of macroinvertebrates, microvertebrates, plant fossils and microfossils (ostracods, foraminifera, charophytes, palynomorphs). In addition, we included data from previous expeditions, particularly those on the dinosaur assemblages.
The environmental model of the Tendaguru Beds presented herein comprises three broad palaeoenvironmental units in a marginal marine setting: (1) Lagoon-like, shallow marine environments above fair weather wave base and with evidence of tides and storms. These formed behind barriers such as ooid bar and siliciclastic sand bar complexes and were generally subject to minor salinity fluctuations. (2) Extended tidal flats and low-relief coastal plains. These include low-energy, brackish coastal lakes and ponds as well as pools and small fluvial channels of coastal plains in which the large dinosaurs were buried. Since these environments apparently were, at best, poorly vegetated, the main feeding grounds of giant sauropods must have been elsewhere. Presumably, tidal flats and coastal plains were visited by dinosaurs primarily during periods of drought. (3) Vegetated hinterland. Vegetation of this environment can only be inferred indirectly from plant material transported into the other depositional environments. Vegetation was dominated by a diverse conifer flora, which apparently formed part of the food source of large herbivorous sauropods. Evidence from various sources suggests a subtropical to tropical palaeoclimate, characterised by seasonal rainfall alternating with a pronounced dry season during the Late Jurassic. In Early Cretaceous times, sedimentological and palaeontological proxies suggest a climatic shift towards more humid conditions.
Die Tendaguru-Schichten von Tansania in Ostafrika (Oberjura bis Unterkreide) sind als Lagerstätte oberjurassischer Dinosaurier seit nahezu einem Jahrhundert weltweit bekannt. Anhand von sedimentologischen und paläontologischen Daten, die während der Deutsch-Tansanischen Tendaguru Expedition 2000 im Typus-Gebiet der Tendaguru-Schichten gewonnen wurden, werden Paläo-Ökosysteme rekonstruiert. Grundlage der Rekonstruktionen sind die Auswertung sedimentologischer Daten sowie die paläo-ökologische Analyse von Makroinvertebraten, Mikrovertebraten, pflanzlichen Fossilien und Mikrofossilien (Ostrakoden, Foraminiferen, Charophyten, Palynomorphen). Darüber hinaus werden Informationen über Dinosaurier berücksichtigt, die bei früheren Expeditionen gewonnen wurden.
Das hier vorgestellte Ablagerungsmodell der Tendaguru-Schichten umfaßt drei Teilbereiche eines randlich marinen Sedimentationsraumes, die wie folgt gekennzeichnet werden können: (1) Lagunen-artige, marine Flachwasserbereiche, die oberhalb der Schönwetter-Wellenbasis lagen und unter deutlichem Einfluß von Gezeiten und Stürmen standen. Sie waren vom offenen Meer durch Barrieren, wie Ooidbarren und siliziklastischen Sandbarrenkomplexen, getrennt und wiesen einen leicht schwankenden Salzgehalt auf. (2) Ausgedehnte Wattgebiete und flache Küstenebenen. Dort befanden sich niedrig-energetische, brackische Strandseen und Teiche sowie Tümpel und kleinere Flußrinnen, in denen die großen Dinosaurier eingebettet wurden. Da diese Lebensräume bestenfalls dürftig bewachsen waren, müssen die Nahrungsquellen und der eigentliche Lebensraum der riesigen Sauropoden anderswo gelegen haben. Vermutlich wurden die Wattgebiete und Flachküsten von Dinosauriern vorrangig in den Trockenzeiten aufgesucht. (3) Bewachsenes Hinterland. Die Vegetation dieses Lebensraumes kann nur indirekt aus Pflanzenresten erschlossen werden, die in die anderen Ablagerungsraume transportiert wurden. Die Vegetation wurde von einer diversen Koniferenflora dominiert, die zumindest teilweise die Nahrungsgrundlage der großen, herbivoren Sauropoden bildete. Sedimentologische und paläontologische Indikatoren sprechen für ein subtropisches bis tropisches Klima wahrend der späten Jurazeit mit einem jahreszeitlichen Wechsel von Regenfällen und ausgeprägten Trockenzeiten. In der frühen Kreidezeit deutet sich ein Wechsel zu starker humiden Bedingungen an.
doi:10.1002/mmng.20020050103
... Na América do Norte, estão representados por Mierasaurus bobyoungi e Moabosaurus utahensis do Cretácico Inferior (Britt et al. 2017;Royo-Torres et al. 2017b). Em África, estão representados por Narindasaurus thevenini e possivelmente Tendaguria tanzaniensis; do Jurássico Médio (Bathoniano) e do Jurássico Superior (Tithoniano), respetivamente (Bonaparte et al. 2000;Mannion et al. 2019;Royo-Torres et al. 2021 Carballido et al. 2011;D'Emic 2012;Mannion et al. 2013Mannion et al. , 2019Mocho et al. 2014;Dai et al. 2022). ...
O registo fóssil de dentes de dinossáurios é abundante, existindo numerosas ocorrências de forma isolada. Alguns autores têm aplicado metodologias de estatística multivariada em dentes de dinossáurios terópodes. Este estudo pretende quantificar, caracterizar e classificar a variabilidade morfológica em dentes de dinossáurios saurópodes através da estatística multivariada, usando a análise de componentes principais (PCA), análise de funções discriminantes (DFA) e análise de clusters, de forma a testar a utilidade destas metodologias para identificar dentes isolados de saurópodes. Os saurópodes constituem um dos grupos de dinossáurios com maior sucesso evolutivo, dominando as faunas de vertebrados do Mesozoico e alcançando uma distribuição geográfica ampla. Eram animais herbívoros e a sua morfologia dentária foi uma característica chave na sua evolução, distinguindo-se no registo fóssil quatro tipos morfológicos principais de dentes: em forma de colher, espátula, cone a cinzel comprimido e lápis. Neste estudo, construiu-se uma base de dados através da bibliografia existente e do registo fotográfico de vários exemplares depositados em museus, com medição indireta de treze variáveis morfométricas. Este trabalho pretende averiguar a utilidade da estatística multivariada para caracterizar os morfo-espaços das morfologias de dentes de saurópodes e identificar as variáveis que melhor descrevem a variabilidade existente; testar a aplicação da estatística multivariada para classificar dentes em forma de colher, de espátula, e de cone a cinzel comprimido; e analisar a potencial influência do desgaste dos dentes nas análise de funções discriminantes (DFA) e na análise de componentes principais (PCA). Os PCAs permitiram identificar algumas variáveis que afetam de forma mais significativa a variabilidade morfológica existente: (i) longitude apicobasal do ápice (LABA); (ii) longitude total da margem distal do ápice (LD); e (iii) longitude total da margem mesial do ápice (LM). Os DFAs permitiram distinguir os diferentes morfo-espaços dos principais grupos analisados (Camarasauridae, Turiasauria, Mamenchisauridae, Somphospondyli e Brachiosauridae). A nível genérico e/ou específico, os morfo-espaços tendem a sobrepor-se e as percentagens de espécimes identificados corretamente diminuem. Não existem diferenças significativas entre os morfo-espaços das distintas espécies de Camarasaurus e os diferentes géneros de Turiasauria. A análise de clusters permitiu agrupar os exemplares de acordo com as suas semelhanças, sendo evidente, por um lado, a agrupação entre dentes espatulados com baixos valores de SI (e.g. Camarasaurus) e os dentes em forma de coração (e.g. Turiasauria) e, por outro, entre os dentes em forma de cone a cinzel comprimido (e.g. Brachiosauridae) e os dentes espatulados com valores de SI moderados (e.g. Mamenchisauridae). Estas análises permitiram diferenciar de forma clara as morfologias de dentes em forma de colher, espátula e cone a cinzel comprimido e caracterizar os seus morfo-espaços, com percentagens de identificação correta superiores a 60% para os principais grupos considerados. O desgaste influencia os resultados obtidos, em particular, faz aumentar a importância das variáveis morfométricas associadas à morfologia do ápice. O registo fóssil de dentes de saurópodes do Jurássico Superior português é relativamente rico em espécimes isolados, atribuíveis a Turiasauria e Macronaria. As análises DFA classificaram quase todos os dentes isolados em forma de coração como Turiasauria, corroborando as classificações prévias. Os dentes espatulados foram atribuídos a distintos taxa, como Turiasauria e Camarasaurus grandis, facto que poderá estar relacionado com o reduzido tamanho da amostra deste tipo de dentes e com a leve sobreposição entre os morfo-espaços de Camarasauridae e Turiasauria. A morfometria multivariada demonstrou ser uma ferramenta com potencial para classificar dentes de saurópodes a um nível supra-genérico.
... The subcircular shape of the proximal articular surface is similar to that of Apatosaurus louisae and distinct to the teardrop shaped of Galeamopus pabsti (Gilmore 1936;Tschopp and Mateus 2017;González Riga et al. 2019) in contrast to the 'D'shaped metacarpals of some titanosaurs as Opisthocoelicaudia (Borsuk-Bialynicka 1977) or MB.R.2093.1-12 (assigned with doubt to Janenschia robusta; Janensch 1961; Bonaparte et al. 2000). This metacarpal is short and robust. ...
... The lateral surface of the proximal end of the right tibia exhibits a well-defined, proximodistally oriented ridge that is present in many titanosauriforms, such as in Dreadnoughtus (Ullmann and Lacovara, 2016: fig. 13a), and has been coined as the "second cnemial crest" (Bonaparte et al., 2000;Mannion et al., 2013). Wiechmann (1999b) also illustrated this crest in the left tibia (Fig. 14B). ...
Dinosaur fossils from the latest Cretaceous (Campanian–Maastrichtian) of Africa and the Arabian Peninsula are rare. Most discoveries to date have consisted of limited fossils that have precluded detailed phylogenetic and paleobiogeographic interpretations. Fortunately, recent discoveries such as the informative Egyptian titanosaurian sauropod dinosaur Mansourasaurus shahinae are beginning to address these long-standing issues. Here we describe an associated partial postcranial skeleton of a new titanosaurian taxon from the Upper Cretaceous (Campanian) Quseir Formation of the Kharga Oasis, Western Desert of Egypt. Consisting of five dorsal vertebrae and 12 appendicular elements, Igai semkhu gen. et sp. nov. constitutes one of the most informative dinosaurs yet recovered from the latest Cretaceous of Afro-Arabia. The relatively gracile limb bones and differences in the coracoid and metatarsal I preclude referral of the new specimen to Mansourasaurus. Both model-based Bayesian tip-dating and parsimony-based phylogenetic analyses support the affinities of Igai semkhu with other Late Cretaceous Afro-Eurasian titanosaurs (e.g., Mansourasaurus, Lirainosaurus astibiae, Opisthocoelicaudia skarzynskii), a conclusion supported by posterior dorsal vertebrae that lack a postzygodiapophyseal lamina, for example. Igai semkhu strengthens the hypothesis that northern Africa and Eurasia shared closely related terrestrial tetrapod faunas at the end of the Cretaceous and further differentiates this fauna from penecontemporaneous assemblages elsewhere in Africa, such as the Galula Formation in Tanzania, that exhibit more traditional Gondwanan assemblages. At present, the specific paleobiogeographic signal appears to vary between different dinosaur groups, suggesting that Afro-Arabian Cretaceous biotas may have experienced evolutionary and paleobiogeographic histories that were more complex than previously appreciated.
... More convincing explanations for the substantial character conflict surrounding the evolutionary position of Bellusaurus and Daanosaurus can be found by considering the issues of taxonomic sampling inherent in this and previous phylogenetic analyses. Several of the features shared by Bellusaurus, Daanosaurus and unambiguous mamenchisauridssuch as anteriorly bifurcated cervical ribs, a bulging or tab-like convexity of the PRDL of middle cervical vertebrae, middle caudal neural spines that approximately double in anteroposterior width towards their summit, and the absence of a 'second cnemial crest' (Bonaparte et al., 2000) on the lateral edge of the proximal tibiahave only recently come to light as a result of concerted comparative study of mamenchisaurids (Mannion et al., 2013(Mannion et al., , 2019aMoore et al., 2020;Upchurch et al., 2021;this study). Numerous taxonomically important and skeletally wellrepresented Middle-Late Jurassic East Asian eusauropods lack comprehensive anatomical descriptions, and ...
The sauropod genus Mamenchisaurus, from the Late Jurassic–Early Cretaceous of East Asia, has a convoluted taxonomic history. Although included in the first cladistic analysis of sauropods, only recently has the monophyly of Mamenchisaurus, and the anatomical diversity of the many penecontemporaneous East Asian eusauropods, been evaluated critically. Here, we re-describe the holotype and only specimen of M. sinocanadorum. Although the original diagnosis is no longer adequate, we identify several autapomorphies that support the validity of this species, including an elongate external mandibular fenestra and distinctive pneumatic structures on the cervical centra. We incorporate new data into a phylogenetic character matrix that also includes Bellusaurus and Daanosaurus, both of which are known only from juvenile material and are often hypothesized to be neosauropods (or close relatives thereof). We recover all species of Mamenchisaurus as part of a radiation of predominantly Middle–Late Jurassic East Asian eusauropods, but the genus is non-monophyletic, underscoring the need for further systematic revision of mamenchisaurid taxonomy. Analyses that score ontogenetically variable characters ambiguously recover Bellusaurus and Daanosaurus as juvenile mamenchisaurids, a hypothesis supported by several features that are unique to mamenchisaurids or exhibit little homoplasy, including anteriorly bifurcate cervical ribs. Finally, computed-tomography reveals extensive vertebral pneumaticity in M. sinocanadorum that is comparable to that of the largest sauropods, and updated scaling analyses imply a neck over 14 m long, rivalling estimates for other exceptionally long-necked sauropods. Previous work has suggested that the elongated cervical ribs of particularly long-necked sauropods such as M. sinocanadorum stabilized the neck by limiting its mobility. Given that extent of pneumaticity responds dynamically to a bone’s habitual loading, we propose that long cervical ribs – and other structural modifications that limited flexibility – promoted the evolution of increasingly long necks by producing a more predictable biomechanical milieu amenable to increased pneumatization.
... At least 12 different species of dinosaurs from the Tendaguru are currently considered to be valid. The largest taxonomic variety of taxa is preserved in sauropods (e.g., Janensch, 1929;Bonaparte et al., 2000;Remes, 2009;Taylor, 2009;Tschopp et al., 2015;Mannion et al., 2019), and re-evaluation of the material, even after more than 100 years of study, has led to the discovery of new taxa (Remes, 2007;Mannion et al., 2019). A high taxonomic diversity also seems to be present in the theropods from the Tendaguru area, although their fossil record is complicated by a dominance of isolated teeth and only few skeletal remains, which makes taxonomic determination in most cases difficult (Rauhut, 2011). ...
... Galeamopus), especially Rebbachisauridae, such as Lavocatisaurus, Zapalasaurus, MMCh-PV-45/34, and MPEF-PV-1698 (Salgado et al. 2006;Allain and Aquesbi 2008;Tschopp et al. 2015;Canudo et al. 2018). In lateral view, the proximal surface of the tibia is convex and posteriorly inclined, as in Dongbeititan, Janenschia, and Lavocatisaurus (Bonaparte et al. 2000;Wang et al. 2007;Mannion et al. 2019). The anterodorsal margin of the proximal epiphysis forms an almost right angle with the lateral surface of the bone ( Figure 14C, E), whereas the posterodorsal margin is more prominent, as seen in the rebbachisaurids Lavocatisaurus and Zapalasaurus, and the derived Titanosauriformes Huabeisaurus, Sauroposeidon, and Uberabatitan (Salgado et al. 2006;Rose 2007;Salgado and Carvalho 2008;D'Emic et al. 2013;Canudo et al. 2018). ...
The Lohan Cura Formation (Albian) at the Cerro de los Leones locality (Neuquén Province, Patagonia, Argentina) yielded several fossil materials, especially sauropod specimens. Among these, Agustinia ligabuei includes postcranial elements of a single individual, with widely debated taxonomy and phylogeny. Here, we provide an extended osteological description and illustrations of the axial and appendicular elements of Agustinia, as well as a revised diagnosis. Moreover, the phylogenetic analysis including a new combination of morphological features recognises Agustinia as a basal Rebbachisauridae, closely related with other South American rebbachisaurids. Our results suggest a more diversified sauropod fauna in the Neuquén Basin, where different members of both neosauropod lineages (i.e. Macronaria and Diplodocoidea) survived in the same region during the Albian age. The reassessment of Agustinia as a basal rebbachisaurid improves our knowledge about the early stages of evolutionary history of Rebbachisauridae, adding new information on the morphological and taxonomic diversification of the clade during the Early Cretaceous of southwestern Gondwana.
... The anterodorsal surface of the pubis is rough and anteriorly extended in lateral view and bears the insertions for the musce ambiens. In Ligabuesaurus, the anterior margin does not form a prominent ambiens process, as seen in Janenschia Wild, 1991(Bonaparte et al., 2000, some brachiosaurids (Giraffatitan and Vouivria Mannion, Allain & Moine, 2017;Janensch, 1961;Mannion et al., 2017) and several flagellicaudantans (Apatosaurus, Dicraeosaurus and Diplodocus; Marsh, 1877;Hatcher, 1901;Janensch, 1929). Posteriorly, a low step divides the iliac peduncle from the acetabular region, which is posteromedially inclined and slightly narrower than the anterior half of the proximal pubis (Fig. 22C), as in Camarasaurus, Tangvayosaurus and Tastavinsaurus (McIntosh et al., 1996;Allain et al., 1999;Canudo et al., 2008). ...
Osteological knowledge of the sauropod dinosaur Ligabuesaurus leanzai is increased by the description of new postcranial elements assigned to the holotype MCF-PVPH-233. Furthermore, a newly referred specimen, MCF-PVPH-228, is recognized after a detailed revision of the abundant sauropod material collected from the Lohan Cura Formation outcrops in the Cerro de los Leones locality (southern Neuquén Basin, Patagonia, Argentina). Recent laboratory preparation and fieldwork allowed us to recognize several new morphological features of the pectoral and pelvic girdles and the cervical and caudal anatomy. Thus, a new diagnosis of Ligabuesaurus is proposed that includes new autapomorphies and a unique combination of features. A phylogenetic analysis based on this new material recovers Ligabuesaurus as a non-titanosaurian somphospondylan, more derived than Sauroposeidon. Therefore, we discuss the palaeobiogeographical implications for the diversification and distribution of South American somphospondylans, especially in the Neuquén Basin, which are closely related to the early stages of evolution of Titanosauria. In this context, Ligabuesaurus represents one of the more complete Early Cretaceous Titanosauriformes and the earliest non-titanosaurian somphospondylan of South America. Finally, the new information on Ligabuesaurus contributes not only to reconstruction of the sauropod faunal composition of south-western Gondwana, but also sheds light on the early stages and emergence of titanosaurians.
... 36), Chuanjiesaurus (Sekiya, 2011:fig. 40), Haestasaurus (Upchurch et al., 2015), Janenschia (Bonaparte et al., 2000;Mannion et al., 2019a), and several titanosaurs (Upchurch, 1995;Wilson and Carrano, 1999;Upchurch et al., 2004a). In Rhomaleopakhus, the posteromedially directed process of the proximal end creates a concavity on the posteromedial surface that does not fade out until approximately the midlength of the element, whereas the lateral surface is flat or slightly convex anteroposteriorly. ...
Hudiesaurus sinojapanorum is a Late Jurassic sauropod from northwestern China that was erected on the basis of a cervicodorsal vertebra, four teeth, and a nearly complete forelimb. However, re-evaluation of this material, and comparisons with other taxa, indicate that there are few grounds for regarding these specimens as congeneric. Consequently, although we retain the vertebra as the holotype specimen of Hudiesaurus, the forelimb is assigned to a new taxon—Rhomaleopakhus turpanensis, gen. et sp. nov. The teeth previously referred to Hudiesaurus are poorly preserved but resemble those of several other ‘core Mamenchisaurus-like taxa’ (CMTs) from East Asia, such as Mamenchisaurus sinocanadorum. Phylogenetic analyses confirm that Hudiesaurus is a CMT and the sister taxon of Xinjiangtitan. Despite some uniquely shared features, their large size, and close geographic provenance, Hudiesaurus and Xinjiangtitan are retained as distinct genera based on their stratigraphic separation and numerous anatomical differences. Rhomaleopakhus is also shown to be a CMT in all analyses, being most closely related to Chuanjiesaurus and Analong. We link the convergent evolution of robust antebrachia and an enlarged olecranon in CMTs, titanosaurs, and some ornithischians (e.g., ceratopsids) to a more flexed orientation of the forearm, an enhanced role for the forelimb in locomotion, and an anterior shift in the whole-body center of mass. CMTs and titanosaurs potentially converged on a feeding strategy in which the ability to increase browse height via bipedal rearing was sacrificed in return for more efficient locomotion that improved travel between patchily distributed food sources.
... The lateral surface is slightly convex with no other surface features visible, whereas the medial surface appears rugose posteriorly, where the fovea ligamentosa inserted. As the tibia lacks an additional articular ridge with the fibula (the second cnemial crest as described in Bonaparte et al., 2000), MUVP 182 is here conservatively referred to Somphspondyli with a possibility that it represents a titanosaurian sauropod (Mannion et al., 2013). Although compressed and partially preserved, MUVP 182 resembles the same elements recovered for Ampelosaurus, Atsinganosaurus, Lirainosaurus, and Lohuecotitan of Late Cretaceous Europe (Le Loeuff, 2005;Díez Díaz et al., 2013, but no meaningful characters are preserved or present to confidently suggest a close association or deviation away from the general titanosaurian condition. ...
Upper Cretaceous dinosaur remains from Afro-Arabia are rare and mainly restricted to pre-Turonian horizons. Consequently, the discovery of new fossils from Upper Cretaceous deposits in the Western Desert, central Egypt is significant because it adds to the meager record of dinosaurs described from this landmass. The oases of Kharga and Dakhla, Western Desert, Egypt, expose the Quseir Formation (Campanian), with the titanosaurian sauropod Mansourasaurus shahinae as the only currently named dinosaur from strata of this age in the entire region. Numerous (∼80) other non-avian dinosaur fossils have also been collected from the Quseir Formation, including material that can be referred to Somphospondyli, Titanosauriformes/Titanosauria, and non-avian Theropoda. Among the discoveries from the Dakhla Oasis are the proximal ends of an associated sauropod tibia and fibula and several sauropod caudal vertebrae representing both sub-adult and adult individuals. Middle caudal vertebrae can be referred to Titanosauria on the basis of procoelous centra and anteriorly displaced neural arches. One caudal vertebra even exhibits pneumatic internal structures rarely observed outside of the Upper Cretaceous South American saltasaurines. Dinosaur fossils recovered from the Kharga Oasis include a partial femur and partial cervical vertebra of a sauropod dinosaur and an isolated proximal fibula of a non-avian theropod dinosaur. Taken together, these findings indicate that Late Cretaceous North African ecosystems supported a diversity of non-avian dinosaurs, some demonstrating affinities with South American forms and others (e.g., Mansourasaurus) with Eurasian groups. Additional exploration of the uppermost Cretaceous units in southern Egypt offers promise for the discovery of important fossils to better characterize Afro-Arabian biotas generally while also providing important perspectives on non-avian dinosaur faunas near the close of the Mesozoic Era.
... This is an unusual orientation for a sauropod and it is here interpreted as an additional autapomorphy of Europasaurus, which is acquired convergently in Rapetosaurus (Curry Rogers 2009). The 'second cnemial crest' originally described for Janenschia (Bonaparte et al. 2000) is not present in Europasaurus. This 'second cnemial crest', or anterior prominence, is present in several sauropods such as Camarasaurus, Giraffatitan, Lusotitan, Tastavinsaurus, Phuwiangosaurus and Rapetosaurus (Mannion et al. 2013). ...
The Late Jurassic was a period of great diversity for sauropod dinosaurs, with different lineages of Neosauropoda flourishing, including several camarasauromorph taxa. Efforts made in recent years have resulted in a great increase in our current knowledge of basal camarasauromorph evolution, including both the detailed description of new and previously poorly known sauropods and expanded phylogenetic analyses. Although most recent phylogenies converge in their results on early camarasauromorph diversification, the phylogenetic position of Europasaurus, from the Late Jurassic of Germany, remains controversial despite the completeness of the material representing this species. Although Europasaurus was recovered as a relatively basal camarasauromorph in all phylogenetic analyses to date, some of them retrieved this taxon in a slightly more derived position, among basal brachiosaurids. Europasaurus is not only one of the most complete camarasauromorphs but also the first unequivocal dwarf that evolved through paedomorphosis, retaining several plesiomorphic characters, especially in the cranium. Cranial and axial material of Europasaurus has been described in detail but the appendicular skeleton has not. The current paper rectifies this by providing detailed descriptions and illustrations of its appendicular skeleton. In addition, an extensive re-evaluation of the systematic position of Europasaurus was done based on the three most substantial data sets published in recent years. These analyses resolved Europasaurus as a basal camarasauromorph in all cases, but brachiosaurid affinities remain plausible, especially considering the heterochronic evolution of the taxon.
... The first is related to the presence of a secondary cnemial crest as in other somphospondylii (e.g. Janenschia robusta Bonaparte et al. 2000;Mannion et al. 2013Mannion et al. , 2019; Dreadnoughtus schrani, Gobititan shenzhouensis, Antarctisaurus wichmannianus, Uberabatitan riberoi and weakly present in Atsinganosaurus velauciensis, Ullmann and Lacovara 2016) and probably in Lohuecotitan pandafilandi, as well as some specimens of Morphotype I at Lo Hueco Páramo et al. 2017a, b). A recent study noted the presence of this feature in several sauropods (Mannion et al. 2013: Ch261). ...
The titanosaurian appendicular skeleton exhibits morphological similarities among different clades and its osteological information is usually less taxonomically meaningful than those of other regions of the skeleton. There is a probable morphological convergence due to morphofunctional similarities between members of different titanosaurian groups. In addition, higher intraspecific variability has hindered the assessment of similar forms such as the Late Cretaceous titanosaurians of the Ibero-Armorican domain. The use of 3D-geometric morphometrics and discriminant analyses on an abundant sample of titanosaurian limb elements from the Lo Hueco site and other titanosaurian taxa has been able to characterize the differences between several sauropod clades with the control of the intraspecific variability. Similar methods with the use of surface analyses of other titanosaurs enabled the recognition of morphological similarities congruent with morphofunctional convergences between one of the lithostrotian morphotypes identified in Lo Hueco (the Morphotype II) and some gracile colossosaurs such as Mendozasaurus neguyelap. In contrast, Morphotype I at Lo Hueco present a more typical titanosaurian morphology, resembling Jainosaurus cf. septentrionalis. The use of discriminant analyses allowed us to distinguish Colossosauria on the basis of limb morphology for the first time. We also observed that Colossosauria and Saltasauridae diverge from a more typical titanosaurian non-autopodial limb skeleton. Our results also suggest a highly different and specialized limb skeleton for the Saltasauridae in comparison with other sampled titanosaurians. The use of surface semilandmarks and discriminant analyses also allowed us to propose several new potential osteological characters for use in phylogenetic analyses through the maximization of differences between the sampled clades.
... There is no 'tuberculum fibularis' on the posterior surface of the cnemial crest, contrasting with several flagellicaudatans (Harris 2007;Tschopp et al. 2015) and some 'basal' macronarians (Mannion et al. 2017). Poor preservation of the proximal end does not allow us to determine whether a 'second cnemial crest' (Bonaparte et al. 2000;Mannion et al. 2013) was present. ...
Dinosaur remains were discovered in the 1860’s in the Kimmeridgian (Late Jurassic) Reuchenette Formation of Moutier, northwestern Switzerland. In the 1920’s, these were identified as a new species of sauropod, Ornithopsis greppini, before being reclassified as a species of Cetiosauriscus (C. greppini), otherwise known from the type species (C. stewarti) from the late Middle Jurassic (Callovian) of the UK. The syntype of “C. greppini” consists of skeletal elements from all body regions, and at least four individuals of different sizes can be distinguished. Here we fully re-describe this material, and re-evaluate its taxonomy and systematic placement. The Moutier locality also yielded a theropod tooth, and fragmentary cranial and vertebral remains of a crocodylomorph, also re-described here. “C.” greppini is a small-sized (not more than 10 m long) non-neosauropod eusauropod. Cetiosauriscus stewarti and “C.” greppini differ from each other in: (1) size; (2) the neural spine morphology and diapophyseal laminae of the anterior caudal vertebrae; (3) the length-to-height proportion in the middle caudal vertebrae; (4) the presence or absence of ridges and crests on the middle caudal centra; and (5) the shape and proportions of the coracoid, humerus, and femur. These anatomical differences, combined with their discrepancy in stratigraphic age, make it unlikely that C. stewarti and “C.” greppini belong to the same genus, as also supported through our phylogenetic analysis. “C.” greppini cannot be assigned to any other contemporaneous sauropod taxon from Europe, but is diagnosed by an autapomorphic rugosity on the posteromedial margin of the humerus, as well as a unique combination of features. As such, we erect the new genus name Amanzia for the Swiss taxon “Ornithopsis” greppini, augmenting the growing diversity of Late Jurassic European sauropods. Our phylogenetic analysis places it outside of Neosauropoda, either as the sister taxon to that clade, or as a member of Turiasauria.
... Nullotitan differs from saltasaurids (sensu Gonzáles-Riga et al., 2019) in having proximal caudals anteroposteriorly short, with posterior articular surfaces slightly convex, and apneumatic centra. These are plesiomorphic traits indicating that Nullotitan is not related with saltasaurids, which usually have caudal vertebrae with dorsoventrally depressed and strongly procoelous centra, and highly pneumatic centra and neural arches (e.g., Powell, 1993;Salgado et al., 1997;Bonaparte et al., 2000;Salgado & Azpelicueta, 2000;Rose, 2007;Taylor, 2009;González Riga et al., 2019). In this regard, the ventral aspect of the caudal vertebrae of Nullotitan is unique in that the proximal and distal caudals show a relatively flat ventral surface of centrum, whereas mid-caudals exhibit a deep longitudinal furrow surrounded by two thickened ridges. ...
The first fossil remains of vertebrates, invertebrates, plants and palynomorphs of the Chorrillo Formation (Austral Basin), about 30km to the SW of the town of El Calafate (Province of Santa Cruz), are described. Fossils include the elasmarian (basal Iguanodontia) Isasicursor santacrucensis gen. et sp. nov., the large titanosaur Nullotitan glaciaris gen. et sp. nov., both large and small Megaraptoridae indet., and fragments of sauropod and theropod eggshells. The list of vertebrates is also composed by the Neognathae Kookne yeutensis gen. et sp. nov., two isolated caudal vertebrae of Mammalia indet., and isolated teeth of a large mosasaur. Remains of fishes, anurans, turtles, and snakes are represented by fragmentary material of low taxonomical value, with the exception of remains belonging to Calyptocephalellidae. On the other hand, a remarkable diversity of terrestrial and freshwater gastropods has been documented, as well as fossil woods and palinological assemblages. The Chorrillo Formation continues south, in the Las Chinas River valley, southern Chile, where it is called Dorotea Formation. Both units share in their lower two thirds abundant materials of titanosaurs, whose remains cease to appear in the upper third, registering only elasmarians (Chorrillo Formation) and hadrosaurs (Dorotea Formation). Above both units there are levels with remains of invertebrates and marine reptiles. It is striking that the dinosaurs of the lower two thirds of the Chorrillo and Dorotea formations are represented by large basal titanosaurs and Megaraptoridae coelurosaurs, being the Saltasaurinae and Aeolosaurinae sauropods and Abelisauridae theropods totally absent. In contrast, these taxa are dominant components in sedimentary units of central and northern Patagonia (e.g., Allen, Los Alamitos, La Colonia formations). Such differences could reflect, in part, a greater antiquity (i.e., late Campanian-early Maastrichtian) for the Chorrillo fossils, or, more probably, different environmental conditions. Thus, knowledge of the biota of the southern tip of Patagonia is expanded, particularly those temporarily close to the K-Pg boundary.
... 31) indicates that it probably would have been subcircular in contour prior to diagenetic deformation. There is a small but distinct fibular tubercle posterior to the cnemial crest, which projects into a short second cnemial crest (sensu Bonaparte et al., 2000;fig. 31A). ...
The Upper Jurassic Morrison Formation of the western United States preserves one of the best-known Mesozoic paleoecosystems worldwide. The formation crops out over an area from New Mexico and Oklahoma to Montana and Utah and encompasses a time span of approximately eight million years. Recent studies indicate a high diversity of gigantic, herbivorous sauropod dinosaurs, but the geographic and temporal distributions of species or even genera of these animals remain poorly understood. In particular, sauropod specimens from northern outcrops of the formation have rarely been studied in detail, and temporal relationships among sites are imprecise. Here, we reassess the taxonomic diversity of the sauropods from a historic Carnegie Museum locality in northern Wyoming. Previous referrals of material to the well-known diplodocid genera Apatosaurus and Diplodocus cannot be confidently confirmed; instead, all these specimens more likely represent elements from the recently recognized Galeamopus. Specimens previously assigned to Camarasaurus and Haplocanthosaurus could not be referred to these genera based on apomorphies, due to a lack of detailed knowledge concerning the genus- and species-level taxonomy of these sauropods. Our findings imply that many referrals of incomplete diplodocid skeletons to Apatosaurus and Diplodocus must be reassessed. These reassessments are particularly important with regard to specimens from northern localities of the Morrison Formation, as it is becoming increasingly evident that diplodocids from this area were distinct from better-known, more southerly taxa. This geographic segregation does not seem to apply to nondiplodocid sauropods; however, these taxa are also in need of systematic revision, which may reveal species-level patterns similar to those observed in Diplodocidae.
... It is a very robust structure that supported the triceps tendon. Posterior to the cnemial crest, there is no sign of a 'second cnemial crest' (Bonaparte et al., 2000). Laterally, between the cnemial crest and the tibial protuberance, there is a small depression extending proximodistally, where M. extensor digitorium communis inserted. ...
Uberabatitan ribeiroi is a Late Cretaceous titanosaur (Dinosauria, Sauropoda) from southeastern Brazil. Here we provide
a detailed revision of all its available specimens, including new elements from the type-locality. One new autopomorphy
is added to diagnosis of the taxon: astragalus with a well-developed anteroposterior crest that mediodistally delimits the tibial articulation. Linear regressions were conducted in an attempt to circumscribe specimens within the type-series, revealing that it is composed of several individuals, with inferred total body lengths varying from 7 to 26 meters. Phylogenetic analyses including U. ribeiroi show that the Brazilian taxon corresponds to a non-saltasaurid lithostrotian titanosaur.
... The astragalus exhibits the wedge morphology observed in other Neosauropoda (Upchurch, 1995(Upchurch, , 1998. It is wider (mediolaterally) than high (proximodistally), as in Janeschia robusta ( Bonaparte et al., 2000;Figs. 6 and 7), Camarasaurus grandis ( Wilson and Sereno, 1998;Fig. ...
... Full-size DOI: 10.7717/peerj.6348/ fig-3 (Bonaparte, Heinrich & Wild, 2000) and, to a lesser extent, Mierasaurus (Royo-Torres et al., 2017a), strongly resemble that of NHMUK 1871. In contrast, the anteriormost dorsal vertebrae of Turiasaurus have dorsoventrally taller neural spines (Royo-Torres, Cobos & Alcalá, 2006;Royo-Torres et al., 2017a). ...
The Jurassic/Cretaceous (J/K) boundary, 145 million years ago, has long been recognised as an extinction event or faunal turnover for sauropod dinosaurs, with many ‘basal’ lineages disappearing. However, recently, a number of ‘extinct’ groups have been recognised in the Early Cretaceous, including diplodocids in Gondwana, and non-titanosauriform macronarians in Laurasia. Turiasauria, a clade of non-neosauropod eusauropods, was originally thought to have been restricted to the Late Jurassic of western Europe. However, its distribution has recently been extended to the Late Jurassic of Tanzania ( Tendaguria tanzaniensis ), as well as to the Early Cretaceous of the USA ( Mierasaurus bobyoungi and Moabosaurus utahensis ), demonstrating the survival of another ‘basal’ clade across the J/K boundary. Teeth from the Middle Jurassic–Early Cretaceous of western Europe and North Africa have also tentatively been attributed to turiasaurs, whilst recent phylogenetic analyses recovered Late Jurassic taxa from Argentina and China as further members of Turiasauria. Here, an anterior dorsal centrum and neural arch (both NHMUK 1871) from the Early Cretaceous Wealden Supergroup of the UK are described for the first time. NHMUK 1871 shares several synapomorphies with Turiasauria, especially the turiasaurs Moabosaurus and Tendaguria , including: (1) a strongly dorsoventrally compressed centrum; (2) the retention of prominent epipophyses; and (3) an extremely low, non-bifid neural spine. NHMUK 1871 therefore represents the first postcranial evidence for Turiasauria from European deposits of Early Cretaceous age. Although turiasaurs show clear heterodont dentition, only broad, characteristically ‘heart’-shaped teeth can currently be attributed to Turiasauria with confidence. As such, several putative turiasaur occurrences based on isolated teeth from Europe, as well as the Middle Jurassic and Early Cretaceous of Africa, cannot be confidently referred to Turiasauria. Unequivocal evidence for turiasaurs is therefore restricted to the late Middle Jurassic–Early Cretaceous of western Europe, the Late Jurassic of Tanzania, and the late Early Cretaceous of the USA, although remains from elsewhere might ultimately demonstrate that the group had a near-global distribution.
... Metatarsal V (Fig. 2) has a widely expanded proximal end, which strongly tapers into a long slender shaft, similar to the brachiosaurids Giraffatitan brancai (Janensch, 1961) and Sonorasaurus (D'Emic, Foreman & Jud, 2016). In Janenschia and Camarasaurus, the expansion is wide too, but it extends further distally along the shaft ( Fig. 7; Bonaparte, Heinrich & Wild, 2000;Tschopp et al., 2015), whereas in many diplodocids, the proximal expansion is similarly developed as in KUVP 129724 ( Fig. 7; Janensch, 1961;Tschopp & Mateus, 2017). The distal articular surface of mt V of KUVP 129724 is only weakly transversely expanded compared to minimum shaft width, which is similar to Camarasaurus, but different from flagellicaudatans (Janensch, 1961;Remes, 2009;Tschopp et al., 2015;Tschopp & Mateus, 2017), see Table S1 and Fig. 7 ...
A set of associated left pedal elements of a sauropod dinosaur from the Upper Jurassic Morrison Formation in Weston County, Wyoming, is described here. Several camarasaurids, a nearly complete small brachiosaur, and a small diplodocid have been found at this locality, but none match the exceptionally large size of the pedal elements. Next to the associated pedal elements, an isolated astragalus, phalanx and ungual were found, which match the large metatarsals in size. The elements cannot be ascribed to diplodocids due to the lack of a ventral process of metatarsal I. Moreover, the morphology of metatarsal V has a broad proximal end, with a long and narrow distal shaft, which differs from Camarasaurus . The size of the material and a medially beveled distal articular surface of metatarsal IV imply an identification as a brachiosaurid. This is the largest pes ever reported from a sauropod dinosaur and represents the first confirmed pedal brachiosaur elements from the Late Jurassic of North America. Furthermore, this brachiosaur material (the pes and the small nearly complete specimen) is the northernmost occurrence of brachiosaurids in the Morrison Formation.
... Although proximally incomplete, it is rounded in lateral view and prominent and antero-laterally projected. The projection posterior to the cnemial crest ('second cnemial crest', sensu Bonaparte et al. 2000) is absent and the cnemial fossa is deep but distally reduced, like in several titanosaurs (Antarctosaurus, Bonitasaura, Rapetosaurus, Opisthocoelicaudia). The medial surface is diagenetically altered in the proximal portion and strongly convex distally, as in all sauropods (Upchurch et al. 2004). ...
Purpose: New sauropod remains (MCF-PVPH-889, MCF-PVPH-899, and MCF-PVPH-900) collected from the Plottier Fm (Coniacian–Santonian) in the south-west of the Neuquén Basin, are here reported. The materials proceed from a fluvial outcrop composed by siltstone and fine sandstone, whose fossil record is known for large-sized sauropod taxa. Methods: Due to the fragmentary condition of the dorsal ribs and the tibia, we focus the description mainly on the femur and the fibula. Results: The specimen MCF-PVPH-889 consists of partially associated postcranial elements represented by a left femur and three fragmentary dorsal ribs of a Titanosauria indet. A right fibula (MCF-PVPH-900) represents another titanosaurian element, while a proximal portion of a right tibia (MCF-PVPH-899) of a smaller individual than others is here referred to as a Sauropoda indet. MCF-PVPH-889 and MCF-PVPH-900 share some features with other titanosaurian taxa (e.g., a femur with medial deflection of the proximal end and elliptical mid-shaft cross-section, a fibula with slightly sigmoidal shaft and well-developed lateral tuberosity) like in Epachthosaurus and Antarctosaurus. However, the femur, with a poorly developed lateral bulge and a relatively low head, and a fibula, with a slender and nearly straight proximal third of the shaft, represent plesiomorphic conditions among titanosaurians. Conclusions: The new remains represent a new sauropod record from the Plottier Fm (Upper Cretaceous). Nevertheless, due to the lack of more diagnostic elements, we prefer to consider the specimens MCF-PVPH-889 and MCF-PVPH-900 as Titanosauria indet, and the MCF-PVPH-899 as Sauropoda indet. This new evidence expands the Coniacian sauropod record of the Neuquén Basin and contributes, in some measure, to our knowledge of the stratigraphical distribution of sauropods from the Patagonian Upper Cretaceous fossil-bearing levels. © 2018, Springer International Publishing AG, part of Springer Nature.
... However, since the type and referred specimens of "Dsungaripterus" brancai are tibiotarsi, more material is necessary to establish whether only one, or two taxa of dsungaripteroids were present in the Tendaguru fauna. The dinosaur fauna from Tendaguru is dominated by sauropods, which are re- presented by at least seven species, the brachiosaurid Brachiosaurus brancai, the probably basal titanosaurs Janenschia ro- busta and Tendaguria tanzaniensis (which might be conspecific), the dicraeosaurids Dicraeosaurus hansemanni and D. sattleri, and the diplodocids Tornieria africana and Australodocus bohetii (Janensch 1914, 1929, Bonaparte et al. 2000, Remes 2006. Theropod di- nosaurs are represented by the basal neo- ceratosaur Elaphrosaurus bambergi ( Fig. 9) and at least six more taxa (Janensch 1920, Rauhut 2005b, c, 2006b). ...
... The two margins end in two pronounced, bulge-like posteroventral expansions. The two expansions are separated by a strongly concave posteroventral margin in ventral view, similar to the condition considered autapomorphic in Janenschia robusta (Bonaparte, Heinrich & Wild, 2000). The tibial fossa is larger than the fibular fossa and subdivided by a shallow, oblique, anteroposteriorly oriented ridge into a medial and a lateral portion. ...
Diplodocids are among the best known sauropod dinosaurs. Numerous specimens of currently 15 accepted species belonging to ten genera have been reported from the Late Jurassic to Early Cretaceous of North and South America, Europe, and Africa. The highest diversity is known from the Upper Jurassic Morrison Formation of the western United States: a recent review recognized 12 valid, named species, and possibly three additional, yet unnamed ones. One of these is herein described in detail and referred to the genus Galeamopus. The holotype specimen of Galeamopus pabsti sp. nov., SMA 0011, is represented by material from all body parts but the tail, and was found at the Howe-Scott Quarry in the northern Bighorn Basin in Wyoming, USA. Autapomorphic features of the new species include a horizontal canal on the maxilla that connects the posterior margin of the preantorbital and the ventral margin of the antorbital fenestrae, a vertical midline groove marking the sagittal nuchal crest, the presence of a large foramen connecting the postzygapophyseal centrodiapophyseal fossa and the spinopostzygapophyseal fossa of mid- and posterior cervical vertebrae, a very robust humerus, a laterally placed, rugose tubercle on the concave proximal portion of the anterior surface of the humerus, a relatively stout radius, the absence of a distinct ambiens process on the pubis, and a distinctly concave posteroventral margin of the ascending process of the astragalus. In addition to the holotype specimen SMA 0011, the skull USNM 2673 can also be referred to Galeamopus pabsti. Histology shows that the type specimen SMA 0011 is sexually mature, although neurocentral closure was not completed at the time of death. Because SMA 0011 has highly pneumatized cervical vertebrae, the development of the lamination appears a more important indicator for individual age than neurocentral fusion patterns. SMA 0011 is one of very few sauropod specimens that preserves the cervico-dorsal transition in both vertebrae and ribs. The association of ribs with their respective vertebrae shows that the transition between cervical and dorsal vertebrae is significantly different in Galeamopus pabsti than in Diplodocus carnegii or Apatosaurus louisae, being represented by a considerable shortening of the centra from the last cervical to the first dorsal vertebra. Diplodocids show a surprisingly high diversity in the Morrison Formation. This can possibly be explained by a combination of geographical and temporal segregation, and niche partitioning.
... In the caudal region, many sauropods retain the ancestral amphicoelous condition (e.g., Limaysaurus, Calvo and Salgado, 1995;Camarasaurus, Osborn and Mook, 1921). (Young, 1954); (2) in the diplodocoid clade Flagellicaudata (Calvo and Salgado, 1995); (3) in the titanosaur clade Lithostrotia (McIntosh, 1990;Wilson and Sereno, 1998;Upchurch et al., 2004); and (4) in a tail of uncertain taxonomic position once referred to the Late Jurassic Janenschia (Bonaparte et al., 2000). In some Late Cretaceous lithostrotians, middle and distal caudal vertebrae are also procoelous ( Fig. 2; McIntosh, 1990;Wilson, 2002;Upchurch et al., 2004). ...
... km north-north-east of Tendaouru Hill . S t r a t i o r a p h y : Upper Saurian Bed Ta x o n : Jruw»schia mbusia NLinlhcr of taxa : 1 D e s i g n a t i o n : Multi-i11dividttal but sin,le-taxon assenlblage 21-22) Four partial skeletons of Janenschia robusta were uncovered, along with additional scattered postcranial bones (GTE field catalo- gue; Janensch 1922Janensch , 1929a Bonaparte et al., in press) . According to Janensch's unpublished excavation sketches and hand-written notes (GTE field catalogue: 21), partial skeleton I consists of a reasonably complete left hind limb, with femur, tibia, fibula, and the foot, including astragalus, metacarpals, and phalanges. ...
Tendaguru is one of the most important dinosaur localities in Africa. The Tendaguru Beds have produced a diverse Late Jurassic (Kimmeridgian to Tithonian) dinosaur assemblage, including sauropods (Brachiosaurus, Barosaurus, Dicraeosaurus, Janenschia), theropods (e.g., Elaphrosaurus, Ceratosaurus, Allosaurus), and ornithischians (Kentrosaurus, Dryosaurus). Contrary to the well studied skeletal anatomy of the Tendaguru dinosaurs, the available taphonomic information is rather limited, and a generally accepted taphonomic model has not yet been established. Assessment of unpublished excavation sketches by the German Tendaguru expedition (1909–1913) document bone assemblages of sauropod and ornithischian dinosaurs from the Middle Saurian Bed, Upper Saurian Bed, and the Transitional Sands above the Trigonia smeei Bed, and shed some light on the taphonomy of the Tendaguru dinosaurs. Stages of disarticulation range from incomplete skeletons to solitary bones, and strongly argue for carcass decay and post-mortem transport prior to burial. The sauropod bone accumulations are dominated by adult individuals, and juveniles are rare or missing. The occurrence of bones in different superimposed dinosaur-bearing horizons indicates that skeletal remains were accumulated over a long time span during the Late Jurassic, and the majority of the bone accumulations are probably attritional. These accumulations are likely to have resulted from long-term bone imput due to normal mortality events caused by starvation, seasonal drought, disease, old age and weakness. The depositional environment of the Middle and Upper Saurian Bed was mainly limnic to brackish in origin, while the palaeoenvironment of the Transitional Sands was marginal marine.
Tendaguru zählt zu den bedeutendsten Dinosaurier-Lagerstätten Afrikas. Aus den Tendaguru-Schichten sind zahlreiche Skelettreste von Sauropoden (Brachiosaurus, Barosaurus, Dicraeosaurus, Janenschia), Theropoden (z.B. Elaphrosaurus, Ceratosaurus, Allosaurus) und Ornithischiern (Kentrosaurus, Dryosaurus) geborgen worden. Sie stammen aus der späten Jura-Zeit (Kimmeridge — Tithon). Während der Skelettbau der Tendagurusaurier gut untersucht ist, wirft die Taphonomie des Sauriervorkommens von Tendaguru noch immer Fragen auf. Unklar ist bislang, wie die enormen Anreicherungen von Dinosaurierknochen in den Tendaguru-Schichten zustandekamen. Unveröffentlichte Grabungsskizzen der Deutschen Tendaguru Expedition (1909–1913) erweitern unsere Kenntnisse über die Taphonomie der Tendagurusaurier. In den ausgewerteten Grabungsskizzen sind Knochenansammlungen von Sauropoden und Ornithischiern aus dem Mittleren und Oberen Sauriermergel sowie aus den Übergangsschichten über der Trigonia smeei-Schicht dokumentiert. Die Lage und der Erhaltungszustand der Funde lassen auf erheblichen Zerfall der Kadaver und post-mortalen Transport von Skelettelementen vor der Einbettung schließen. Das Vorkommen von Saurierknochen in mehreren übereinanderliegenden Profilabschnitten der Tendaguru-Schichten zeigt, daß Skelettreste während der späten Jura-Zeit über einen längeren Zeitraum hinweg akkumuliert wurden. Die Ansammlungen von Skelettresten gehen wahrscheinlich auf „normale” Sterbe-Ereignisse zurück, wie z. B. Verhungern, Verdursten, Kankheit, Altersschwäche und jahreszeitliche Dürre. Als Ablagerungsraum der Mittleren und Oberen Saurierschicht kommt ein küstennaher limnischer, zeitweise wohl auch brackischer Küstenstreifen in Betracht. Die knochenführenden Übergangsschichten unter- und oberhalb der Saurierschichten sind randlich marine Ablagerungen.
doi:10.1002/mmng.1999.4860020102
... The comparison with other figured caudal series (e.g . Janensch 1950;Osborn & Mook 1921;Bonaparte et al. 2000) suggests that CdH (MG 4985-9) should be a caudal vertebra located around the fifteenth position. ...
Lusotitan atalaiensis was one of the first sauropod taxa established for the Upper Jurassic of the Lusitanian Basin (Portugal). The lectotype of L. atalaiensisfound in Peralta (Lourinhã) was firstly considered as new species of Brachiosaurus, considering the similarities with Brachiosaurusand Giraffatitan. This specimen was originally considered as a brachiosaurid, and more recently, a cladistics analysis, suggested Lusotitan as a basal macronarian and a possible member of Brachiosauridae. In this study, new information is provided about the Lusotitanlectotype, with reinterpretation and description of previously described and undescribed elements (e.g. sacral vertebrae, chevrons, ulna, ischium, pubis or fibula). The validity of this taxon is confirmed by a revised diagnosis. The present comparative study and the phylogenetic analyses suggest the assignation of Lusotitanto Brachiosauridae supported by the presence of pronounced and dorsoventrally short deltopectoral crest, short ischiatic contribution to the acetabulum and radius length/tibia length ratio >1. The presence of brachiosaurids in the Portuguese Upper Jurassic record is confirmed, supporting the presence of a wide paleobiogeographic distribution for Brachiosauridae during the Late Jurassic, being present in Europe, North America and Africa
... The comparison with other figured caudal series (e.g . Janensch 1950;Osborn & Mook 1921;Bonaparte et al. 2000) suggests that CdH (MG 4985-9) should be a caudal vertebra located around the fifteenth position. ...
Mocho P, Royo-Torres R, Ortega F. 2016. Evolutionary history of Late Jurassic sauropods of the Lusitanian Basin (Portugal) and presentation of a new basal macronarian form, p.190. In: Farke A, MacKenzie A, Miller-Camp J (eds.), Society of Vertebrate Paleontology, October 2016, Abstract of papers, 76th Annual Meeting, Grand America Hotel, Salt Lake City, Utah, USA, October 26–29, 2016
... With an RI of 0.28, this element is considered gracile using the metric of Wilson and Upchurch (2003) and categories of Salgado et al. (2014;RI >0.3 considered robust). A large, lateral bulge (second cnemial crest of Bonaparte et al., 2000) forms the cranial portion of the proximal tibial articular surface (Fig. 13E). The cnemial crest originates near the midpoint of the medial face in proximal view. ...
The postcranial anatomy of giant titanosaurians remains poorly known because of a combination of preservational and collection biases. Dreadnoughtus schrani, a recently described, large titanosaur from the Campanian–Maastrichtian Cerro Fortaleza Formation of Santa Cruz Province, Argentina, offers the first opportunity for detailed study of appendicular anatomy of a truly giant titanosaurian. The entire appendicular skeleton is represented except the manus and portions of the pes. Comparisons with related titanosauriforms reveal that the holotype skeleton (MPM-PV 1156) exhibits three appendicular autapomorphies: (1) a cranioventrally-caudodorsally oriented ridge across the medial surface of the cranial end of the scapular blade; (2) a distinct concavity on the caudomedial surface of the proximal radius; and (3) the distal end of the radius is subrectangular with subequal craniocaudal and mediolateral dimensions. Appendicular synapomorphies shared between Dreadnoughtus and other titanosauriforms encompass a wide range of body sizes, from the giant Argentinosaurus to the dwarf Magyarosaurus. Among described titanosauriforms, only a single feature occurs exclusively among the appendicular skeletons of the largest taxa: an accessory ventrolateral process is present on the preacetabular lobe of the ilium in Dreadnoughtus, Alamosaurus, and Giraffatitan. This process appears to have arisen in response to greater stress applied by hind limb adductor musculature in these giant terrestrial vertebrates. Continued investigation of titanosaurian anatomy, myology, and biomechanics is needed to gain greater understanding of the functional nature of wide-gauge posture.
SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP
Citation for this article: Ullmann, P. V., and K. J. Lacovara. 2016. Appendicular osteology of Dreadnoughtus schrani, a giant titanosaurian (Sauropoda, Titanosauria) from the Upper Cretaceous of Patagonia, Argentina. Journal of Vertebrate Paleontology, e1225303. DOI: 10.1080/02724634.2016.1225303.
... Procoelous anterior caudal vertebrae ( Fig. 1.3) evolved in- dependently three or four times, according to our current understanding of sauropod interrelationships ( (McIntosh, 1990;Wilson and Sereno, 1998;Upchurch et al., 2004); and (4) in a tail of uncertain taxonomic position once referred to the Late Jurassic Janenschia ( Bonaparte et al., 2000). In some Late Cretaceous lithostrotians, middle and distal caudal vertebrae are also procoelous ( Fig. 2; McIntosh, 1990;Wilson, 2002;Upchurch et al., 2004). ...
Sauropod dinosaurs achieved the largest body sizes and the most elongate necks and tails of any terrestrial vertebrate. The elongate, cantilevered necks of sauropods comprised opisthocoelous vertebrae joined at concavo-convex joints. Opisthocoelous centra also occurred in the dorsal region of sauropods and procoelous centra in the tails of certain lineages. Concavo-convex intercentral joints have been hypothesized to increase the flexibility of the vertebral column or to stabilize intervertebral joints against shear stresses. Using Alligator as an extant analog, condyle convexity and range of motion were measured at every intervertebral joint in an individual, with the latter measured in situ. Results reveal that convexity is greatest in the alligator presacral column where flexibility is low; amphiplatyan vertebrae occur in the distal caudal region where flexibility is highest. The negative relationship between convexity and flexibility is not significant, indicating that flexibility is independent of centrum articular morphology. Convexity is greatest in regions in which high shear stresses are predicted to result from terrestrial locomotion and tail flexion. The evolution of opisthocoelous cervical vertebrae in early sauropods likely strengthened the long and massive neck against catastrophic dislocations without compromising joint mobility. The stabilization provided by dorsal opisthocoely and caudal procoely may relate to clade-specific specializations such as the "whiplash" tails of flagellicaudatans and the "wide-gauge" limb stance in titanosaurs. The study of opisthocoely and procoely provides a means to understand the loading regimes of the vertebral column in sauropods and other vertebrates, with implications for the behavior and ecology of fossil taxa.
A new somphospondylan titanosauriform from the Lower Cretaceous of Spain is described from the remains found at the Sant Antoni de la
Vespa site (upper Barremian Arcillas de Morella Formation) located in Morella. Garumbatitan morellensis gen. et sp. nov. is diagnosed by 11
autapomorphies and eight local autapomorphies; and our phylogenetic analyses suggest that Garumbatitan morellensis might correspond to an
early-branching somphospondylan. The presence of several somphospondylan traits in Garumbatitan morellensis supports the somphospondylan
hypothesis. The phylogenetic distribution of some titanosauriform and somphospondylan novelties in the femur (markedly developed lateral
bulge, high shaft eccentricity, linea intermuscularis cranialis, and trochanteric shelf) is discussed. The tarsus and pes of Garumbatitan morellensis
are distinctive, being characterized by the loss of the calcaneum, relative slenderness of the metatarsals II, III, and IV when compared to the
retracted metatarsals I and V, three pedal phalanges in digit IV, and reduced ungual III. The sauropod fauna of the Iberian Peninsula during
the Hauterivian–Aptian shows a complex phylogenetic mosaic, including forms with Laurasian affinities, mainly titanosauriforms (Soriatitan,
Garumbatitan, and possibly Tastavinsaurus and Europatitan), and Gondwanan affinities, the rebbachisaurid Demandasaurus. Faunal exchange
during the Early Cretaceous between the Europe, North America, East Asia, and Africa is plausible.
Neosauropods were the dominant sauropod clade with a global distribution as early as the Late Jurassic. However, its distribution and biogeography in the Middle Jurassic are unclear due to the paucity of phylogenetic evidence for neosauropod taxa of this age. In China, the only reported Middle Jurassic neosauropod, the diplodocoid, has challenged the traditional East Asian Isolation Hypothesis for dinosaur paleobiogeography. Here, based on phylogenetic analysis including Dashanpusaurus dongi from the early Middle Jurassic of southwest China, we demonstrate that this taxon represents the earliest diverging macronarian as well as the stratigraphically lowest neosauropod globally. Our biogeographic analysis together with other geological evidence further indicates that neosauropods achieved a global distribution at least in the early Middle Jurassic while Pangaea was still a coherent landmass.
The vertebrae of sauropod dinosaurs are characterized by complex architecture involving laminae, fossae, and internal chambers of various shapes and sizes. These structures are interpreted as osteological correlates of an intricate system of air sacs and pneumatic diverticula similar to that of birds. In basal sauropods pneumatic features are limited to fossae. Camerae and camellae are internalized pneumatic chambers independently acquired in neosauropods and some Chinese forms. The polycamerate and camellate vertebrae of higher neosauropods are characterized by internal pneumatic chambers of considerable complexity. The independent acquisition of these derived morphologies in Mamenchisaurus, derived diplodocids, and most titanosauriforms is correlated with increasing size and neck length.
The presacrai vertebrae of basal sauropods were probably pneumatized by diverticula of cervical air sacs similar to those of birds. Although pneumatic characters in sauropods are most extensive and complex in presacrai vertebrae, the sacrum was also pneumatized in most neosauropods. Pneumatization of the proximal caudal vertebrae was achieved independently in diplodocids and titanosaurids. In birds, the synsacrum is pneumatized via abdominal air sacs which function primarily in lung ventilation. The presence of pneumatized sacral and caudal vertebrae in neosauropods indicates that abdominal air sacs were probably present in at least some sauropods.
The Late Jurassic Tendaguru Formation of Tanzania, southeastern Africa, records a rich sauropod fauna, including the diplodocoids Dicraeosaurus and Tornieria, and the brachiosaurid titanosauriform Giraffatitan. However, the taxonomic affinities of other sympatric sauropod taxa are poorly understood. Here, we critically reassess and redescribe these problematic taxa, and present the largest phylogenetic analysis for sauropods (117 taxa scored for 542 characters) to explore their placement in Eusauropoda. Janenschia robusta has played a prominent role in discussions of titanosaur origins, with various authors referring at least some remains to Titanosauria, a clade otherwise known only from the Cretaceous. Redescription of the holotype of Janenschia, and all referable remains, supports its validity and placement as a nonneosauropod eusauropod. It forms a clade with Haestasaurus from the earliest Cretaceous of the UK, and the Middle/Late Jurassic Chinese sauropod Bellusaurus. Phylogenetic analysis and CT scans of the internal pneumatic tissue structure of Australodocus bohetii tentatively support a non-titanosaurian somphospondylan identification, making it the only known pre-Cretaceous representative of that clade. New information on the internal pneumatic tissue structure of the dorsal vertebrae of the enigmatic Tendaguria tanzaniensis, coupled with a full redescription, results in its novel placement as a turiasaur. Tendaguria is the sister taxon of Moabosaurus, from the Early Cretaceous of North America, and is the first turiasaur recognized from Gondwana. A previously referred caudal sequence cannot be assigned to Janenschia and displays several features that indicate a close relationship with Middle–Late Jurassic East Asian mamenchisaurids. It can be diagnosed by six autapomorphies, so we erect the new taxon Wamweracaudia keranjei gen. et sp. nov. The presence of a mamenchisaurid in the Late Jurassic of southern Gondwana indicates an earlier and more widespread diversification of this clade than previously realized, prior to the geographic isolation of East Asia. Our revised phylogenetic dataset sheds light on the evolutionary history of Eusauropoda, including supporting a basal diplodocoid placement for Haplocanthosaurus, and elucidating the interrelationships of rebbachisaurids. The Tendaguru Formation shares representatives of nearly all sauropod lineages with Middle Jurassic–earliest Cretaceous global faunas, but displays a greater range of diversity than any of those faunas considered individually. Biogeographic analysis indicates that the Tendaguru sauropod fauna was assembled as a result of three main phenomena during the late Early and/or Middle Jurassic: (1) invasions from Euramerica (brachiosaurids, turiasaurs); (2) endemism in west Gondwana (dicraeosaurids, diplodocids); and (3) regional extinctions that restricted the ranges of once widespread groups (mamenchisaurids, the Janenschia lineage). Multiple dispersals across the Central Gondwanan Desert are required to explain the distributions of Jurassic sauropods, suggesting that this geographic feature was at most a filter barrier that became easier to cross during the late Middle Jurassic.
The titanosaurian sauropod dinosaur Mendozasaurus neguyelap is represented by several partial skeletons from a single locality within the Coniacian (lower Upper Cretaceous) Sierra Barrosa Formation in the south of Mendoza Province, northern Neuquén Basin, Argentina. A detailed revision of Mendozasaurus, including previously undocumented remains from the holotype site, allows us to more firmly establish its position within Titanosauria, as well as enabling an emended diagnosis of this taxon. Autapomorphies include: (1) middle and posterior cervical vertebrae with tall and transversely expanded neural spines that are wider than the centra, formed laterally by spinodiapophyseal laminae that are not connected with the pre- or postzygapophyses; (2) anterior caudal vertebrae (excluding anteriormost) with ventrolateral ridge-like expansion of prezygapophyses; and (3) humerus with divided lateral distal condyle on anterior surface. New remains demonstrate that the presacral vertebrae of Mendozasaurus were not unusually short anteroposteriorly, with this compression instead resulting from taphonomic crushing. Comparative studies of articulated pedes of other taxa allow us to interpret that the pedal formula of Mendozasaurus was 2-2-2-2-0, based on disarticulated bones that form a right hind foot. Mendozasaurus was incorporated into an expanded version of a titanosauriform-focussed phylogenetic data matrix, along with several other contemporaneous South American titanosaurs. The resultant data matrix comprises 84 taxa scored for 423 characters, and our phylogenetic analysis recovers Mendozasaurus as the most basal member of a diverse Lognkosauria, including Futalognkosaurus and the gigantic titanosaurs Argentinosaurus, Notocolossus, Patagotitan and Puertasaurus. Lognkosauria forms a clade with Rinconsauria (Muyelensaurus + Rinconsaurus), with Epachthosaurus and Pitekunsaurus recovered at the base of this grouping. A basal lithostrotian position for this South American clade is well supported, contrasting with some analyses that have placed these taxa outside of Lithostrotia or closer to Saltasauridae. The sister clade to this South American group is composed of an array of near-global taxa and supports the hypothesis that most titanosaurian clades were widespread by the Early-middle Cretaceous.
available for this article.
doi:10.1002/mmng.200600015
The dinosaur fauna of the Kadzi Formation, central Zambezi Valley, Zimbabwe, includes all 4 sauropod genera found at Tendaguru, Tanzania-Barosaurus, Brachiosaurus, Dicreaeosaurus, and Tornieria. Two genera, Barosaurus and Brachiosaurus, are also typical of the Morrison Formation of the W United States. Camarasaurus might also be present at Kadzi. This genus is not known from Tendaguru, but a camarasaurid (Algoasaurus) is found in beds of roughly comparable age on the SE coast of South Africa. The coexistence of Brachiosaurus and Tornieria in the Kadzi Formation suggests that this dinosaur bed was deposited some time between the Middle and Upper Saurian Beds of Tendaguru. The Gokwe Formation (SW Zambezi Valley, Zimbabwe), the Dinosaur Beds of NW Malawi, and the possibly homotaxial fossiliferous post-Karoo pebble beds of the Luangwa Valley in Zambia are probably all of similar age.-from Authors