The influence of caudofemoral musculature on the titanosaurian (Saurischia: Sauropoda) tail skeleton: morphological and phylogenetic implications

Due to their abundance, taxonomic and morphological diversity, wide range of body sizes and broad geographical distribution, titanosaurian sauropods were one of the most important Cretaceous herbivorous dinosaur groups. Consequently, titanosaurs constitute one of the best samples in which to evaluate the relationship between bony structures and unpreserved soft-tissues within Sauropoda. We reconstruct the morphology and interpret the implications of selected soft-tissues associated with the titanosaurian caudal skeleton. These tissues, especially the M. caudofemoralis longus (CFL), exerted a considerable influence on the anatomy of the caudal vertebrae and haemal arches. In all studied titanosaurian taxa, the reconstructed caudofemoral musculature corresponds to one of three principal morphotypes that accord with previously recognised phylogenetic patterns within the clade. Basal titanosaurians had an elongate M. CFL that extended for much of the proximal half of the tail; in saltasaurines, this muscle was much shorter. Non-saltasaurine lithostrotians exhibited an intermediate condition. Furthermore, the differing position of the fourth trochanter, and therefore, the insertion of the caudofemoral muscles, among various titanosaurian taxa suggests distinctions in the locomotor function of these animals.     

Dentition and histology in titanosaurian dinosaur embryos from Upper Cretaceous of Patagonia,Argentina

Abstract: Exceptionally preserved sauropod embryos from the Late Cretaceous Anacleto Formation in Auca Mahuevo (Neuquén Province, Argentina) have provided fundamental information on titanosaurian ontogeny. This paper describes the dental composition, disposition and microstructure of the specimens. Embryonic teeth show size disparity, with lengths that vary from 1 to 3 mm and diameters ranging from 0.15 to 0.26 mm, with the most frequent length values between 2.5 and 3 mm. Apparently, a typical ‘pencil‐like’ tooth morphology and a dental formula of Pm 4, M 7–8/D10? remained constant during titanosaurian ontogeny, whereas the arrangement of teeth in the skull shows notable ontogenetic changes. Absence of wear facets on teeth suggests a lack of prenatal chewing movements. The enamel proportion is significantly higher in embryos than in mature titanosaurs, which suggests that this relationship varies during ontogeny. Embryonic bony tissue is composed of highly vascularized, cellular woven bone. The absence of osteonal tissue, the high degree of vascularization, the presence of numerous osteocytes and poor development of periosteal bone reveals that the Auca Mahuevo titanosaurs would have had a high early growth rate and that they were buried at a relatively advanced embryonic stage.     

Juvenile and adult teeth of the titanosaurian dinosaur Lirainosaurus (Sauropoda) from the Late Cretaceous of Iberia

A large sample of more than 100 teeth of the titanosaurian sauropod Lirainosaurus astibiae from the Late Cretaceous Laño quarry (Northern Spain) has been studied. Most of the teeth are small (crown height less than 13 mm), cylindrical, with parallel edges and smooth enamel; a few larger teeth are tapered and have more ornamented enamel. These differences are regarded here as ontogenetic changes, the small teeth being interpreted as those of juveniles and the large ones as those of subadult or adult individuals. The juvenile teeth also present some differences in the apex of the crown: some of them have a tapered tip, and others have apical and/or mesial/distal facets. The first are probably unerupted or non-functional teeth, while the ones with wear facets are functional teeth. This is the first time a change in the microwear structures of the apical wear facets between juvenile and adult teeth is observed in a titanosaurian taxon. Moreover, this let us to hypothesize a switch in the diet and food processing between the juvenile and adult individuals of Lirainosaurus. Finally, the teeth of Lirainosaurus are different from those of the Late Cretaceous European titanosaurs described to date and also differ from the teeth of basal titanosauriforms from the Early Cretaceous of the Iberian Peninsula.     

Turiasauria-like teeth from the Upper Jurassic of the Lusitanian Basin,Portugal

Turiasauria is a clade of eusauropods with a wide stratigraphic range that could extend from the Bathonian to the lower Aptian including Turiasaurus, Losillasaurus, Zby and putatively, Galveosaurus, Atlasaurus and isolated remains from Middle Jurassic-to-Lower Cretaceous. Some are characterised by the presence of heart-shaped teeth. Several tooth occurrences from the Portuguese Upper Jurassic with this type of morphology (SI: 1.1–1.8) are reported and discussed. If this morphology is regarded as synapomorphic of Turiasauria, the teeth will be tentatively related to this clade. From a sample of 43 teeth, three main morphotypes are described. Three hypotheses might explain the morphological variation: (1) the range of tooth morphologies indicates variation in the jaw, (2) the range of tooth morphologies indicates taxonomic variation or (3) a combination of both. The general wear pattern in morphotypes I and II starts with a distal facet, then the appearance of mesial/apical facet and finally a ‘V’-shaped facet. In morphotype III, the wear begins with a mesial facet. The variability observed for Portuguese Upper Jurassic specimens is congruent with the morphological variability along the tooth row shown by other sauropods with spatulate/spoon-shaped teeth and it is considered the most parsimonious hypothesis to explain it.     

中国广西晚白垩世一新的巨龙类恐龙

记述了采自广西南宁市郊晚白垩世地层中一巨龙类恐龙新属种:右江清秀龙(Qingxiu- saurus youjiangensis gen.et sp.nov.)。新属种正型标本包括以下不关联的头后骨骼:一段较完整的前部尾椎神经棘、左右胸骨板、左右肱骨。它以以下独特特征组合区别于其他巨龙类:前部尾椎神经棘板状结构不发育、相对较高并呈桨状;胸骨与肱骨最大长之比值较低(约0.65)。广西发现的新属种以及近年来报道的巨龙类恐龙材料表明,亚洲巨龙类恐龙的头后骨骼形态变异度高,白垩纪时期这类恐龙曾在亚洲广为分布。     

Enigmatic teeth from the Jurassic–Cretaceous transition of Morocco: The latest known non-mammaliaform cynodonts (Synapsida,Cynodontia) from Africa?

Two tooth morphotypes corresponding to one or two tetrapod species from the Late Jurassic or Earliest Cretaceous locality of Ksar Metlili (KM), Anoual Syncline (eastern Morocco), are reported and described. These teeth cannot be related to any of the identified vertebrate major groups of this site. They are tricusped and uniradiculate, with a high and large main central cusp mesio-distally surrounded by two smaller accessory cusps. Their morphology is reminiscent of several taxa such as pterosaurs, notosuchians and mammals, with which they are compared here. These morphotypes are tentatively referred to cf. Cynodontia indet. They would be the most recent non-mammaliaform cynodonts reported in Africa and among the latest described. The KM specimens display remarkable plesiomorphic dental features with respect to known contemporaneous non-mammaliaform cynodonts. They might indicate the survival of a relict lineage in a North African refugium.     

Using Dental Enamel Wrinkling to Define Sauropod Tooth Morphotypes from the Ca?adón Asfalto Formation,Patagonia, Argentina

The early Middle Jurassic is regarded as the period when sauropods diversified and became major components of the terrestrial ecosystems. Not many sites yield sauropod material of this time; however, both cranial and postcranial material of eusauropods have been found in the Cañadón Asfalto Formation (latest Early Jurassic–early Middle Jurassic) in Central Patagonia (Argentina), which may help to shed light on the early evolution of eusauropods. These eusauropod remains include teeth associated with cranial and mandibular material as well as isolated teeth found at different localities. In this study, an assemblage of sauropod teeth from the Cañadón Asfalto Formation found in four different localities in the area of Cerro Condor (Chubut, Argentina) is used as a mean of assessing sauropod species diversity at these sites. By using dental enamel wrinkling, primarily based on the shape and orientation of grooves and crests of this wrinkling, we define and describe three different morphotypes. With the exception of one taxon, for which no cranial material is currently known, these morphotypes match the local eusauropod diversity as assessed based on postcranial material. Morphotype I is tentatively assigned to Patagosaurus, whereas morphotypes II and III correspond to new taxa, which are also distinguished by associated postcranial material. This study thus shows that enamel wrinkling can be used as a tool in assessing sauropod diversity.     

On the spinosaurid theropod teeth from the early Barremian (Early Cretaceous) Blesa Formation (Spain)

Spinosaurids are one of the most abundant theropods from the Early Cretaceous of the Iberian Peninsula. The published fossil record consists mainly of teeth. A good example is La Cantalera-1 site in the Blesa Formation (Spain). This site represents a marshy environment with periodic droughts, resulting in a non-permanent body of water. Theropod dinosaur teeth are frequent. Seventeen spinosaurid teeth have been studied using qualitative and quantitative features, including a statistical analysis, in order to compare them with the known spinosaurid record from Spain. In La Cantalera-1 we recognise two different morphotypes: Baryonychinae indet. and Spinosaurinae? indet. Baryonychinae indet. was produced by baryonychines close to Baryonyx; teeth with similar morphology have been described in other sites from the Early Cretaceous of the Iberian Peninsula. Teeth tentatively attributed to Spinosaurinae? indet. were produced by spinosaurines close to Spinosaurus. This morphotype could provide new evidence of the presence of spinosaurines in the Early Cretaceous of the Iberian Peninsula. The small size of the teeth from La Cantalera-1 when compared with those from other sites is interpreted as the result of the conditions characteristic of the deposit area, which represents an ecosystem unable to sustain large-sized organisms.     

Sexual differentiation and mate recognition. The dynamics of structure

Sexual differentiation results in an array of distinct sexual morphotypes and two biological sexes are not sufficient to describe all the forms that are observed. It is also plainly impossible to discuss mate recognition without reference to sexual morphology, because the body and the brain are inseparably linked throughout the sexual differentiation process. It follows that we must rework conceptually inadequate notions about sexual morphology before general theories of sexual behaviour can be developed.     

A complete skull of an early cretaceous sauropod and the evolution of advanced titanosaurians

Advanced titanosaurian sauropods, such as nemegtosaurids and saltasaurids, were diverse and one of the most important groups of herbivores in the terrestrial biotas of the Late Cretaceous. However, little is known about their rise and diversification prior to the Late Cretaceous. Furthermore, the evolution of their highly-modified skull anatomy has been largely hindered by the scarcity of well-preserved cranial remains. A new sauropod dinosaur from the Early Cretaceous of Brazil represents the earliest advanced titanosaurian known to date, demonstrating that the initial diversification of advanced titanosaurians was well under way at least 30 million years before their known radiation in the latest Cretaceous. The new taxon also preserves the most complete skull among titanosaurians, further revealing that their low and elongated diplodocid-like skull morphology appeared much earlier than previously thought.     

Variations in dinoflagellate cyst morphology under conditions of changing salinity during the last 2000 years in the Limfjord, Denmark

Morphological variations are examined in the dinoflagellate cysts Spiniferites spp., Lingulodinium polyedrum and Protoceratium reticulatum (=Operculodinium centrocarpum) from a core taken in the Bj?rnsholm Bay, the Limfjord, Denmark. The fjord has a history of changing salinity, and unusual cyst morphotypes are found in the greatest numbers during periods of inferred low salinity. Variation occurs primarily in cyst process morphology, and the aberrant morphotypes have processes that are shorter, thicker and/or more membranous. The different morphotypes are described and compared with other varieties and forms of the three taxa and to other closely related taxa.     

A New Titanosaurian Braincase from the Cretaceous “Lo Hueco” Locality in Spain Sheds Light on Neuroanatomical Evolution within Titanosauria

Despite continuous improvements, our knowledge of the neurocranial anatomy of sauropod dinosaurs as a whole is still poor, which is especially true for titanosaurians even though their postcranial remains are common in many Upper Cretaceous sites worldwide. Here we describe a braincase from the uppermost Cretaceous locality of ‘‘Lo Hueco” in Spain that is one of the most complete titanosaurian braincases found so far in Europe. Although the titanosaurian Ampelosaurus sp. is known from the same locality, this specimen is clearly a distinct taxon and presents a number of occipital characters found in Antarctosaurus and Jainosaurus, which are approximately coeval taxa from southern Gondwana. The specimen was subjected to X-ray computed tomographic (CT) scanning, allowing the generation of 3D renderings of the endocranial cavity enclosing the brain, cranial nerves, and blood vessels, as well as the labyrinth of the inner ear. These findings add considerable knowledge to the field of sauropod paleoneuroanatomy in general and titanosaurian endocast diversity in particular. Compared with that of many sauropodomorphs, the endocast appears only slightly flexed in lateral view and bears similarities (e.g., reduction of the rostral dural expansion) with Gondwanan titanosaurians such as Jainosaurus, Bonatitan, and Antarctosaurus. The vestibular system of the inner ear is somewhat contracted (i.e., the radius of the semicircular canals is small), but less so than expected in derived titanosaurians. However, as far as the new specimen and Jainosaurus can be contrasted, and with the necessary caution due to the small sample of comparative data currently available, the two taxa appear more similar to one another in endocast morphology than to other titanosaurians. Recent phylogenetic analyses of titanosaurians have not included virtually any of the taxa under consideration here, and thus the phylogenetic position of the new Spanish titanosaurian—even its generic, let alone specific, identification—is not possible at the moment. Nevertheless, both the braincase osteology and the endocast morphology suggest that the specimen represents a derived titanosaurian that presumably branched further from the base of Lithostrotia, potentially even near Saltasauridae, comparable in evolutionary terms with Jainosaurus.     

Bacterial community in the tunic matrix of a colonial ascidian<Emphasis Type="Italic"> Diplosoma migrans</Emphasis>

This paper provides the first information on the morphology of different morphotypes of bacteria in the tunic matrix of the colonial ascidian Diplosoma migrans. Ascidians were collected from waters near Helgoland (German Bight, North Sea). The dominant type is represented by extremely high numbers of long, needle-like rods (length 10–30 µm, width 0.5 µm). The bacteria are motile by means of bipolar monotrichous flagella, generating swift sigmoidal movement. Bacteria are already present during different embryonic stages. It is assumed that they are transferred during sexual propagation from the parental colony to its offspring. As a second morphotype, the tunic harbors screw-like bacteria in low numbers (length 4–10 µm, width 0.5 µm). Besides these conspicuous morphotypes, occasionally motile rods with spore-like globules at one end and additional coccoid forms in large quantities of unknown meaning (possibly spores) were found. The taxonomic status and ecological functions of these differently shaped bacterial groups are unclear.Communicated by H.-D. Franke     

Late Triassic dinosaur teeth from southern Belgium

Isolated Dinosaur teeth have been discovered in the Upper Triassic locality of Habay-la-Vieille, in southern Belgium. Ornithischia are represented by three dental morphotypes; two of them closely resemble isolated teeth from the Middle or Upper Jurassic of Portugal and England. The presence of sauropods in the Upper Triassic of Europe is confirmed. Sauropods already had a wide geographical distribution during the Latest Triassic, as fossils have been discovered in South Africa, Thailand and western Europe. At Habay-la-Vieille, sauropods and prosauropods co-existed at the end of the Triassic. Two dental morphotypes may tentatively be referred to as theropod dinosaurs. The study of isolated teeth indicates that dinosaurs were already well diversified in the Latest Triassic of western Europe. To cite this article: P. Godefroit, F. Knoll, C. R. Palevol 2 (2003) 3–11.     

RAPD Analysis and Taxonomic Reconsideration of the Ceropegia aristolochioides Complex (Apocynaceae-Ceropegieae)

Abstract: Flower morphology, karyology and taxonomy of the polymorphic Ceropegia aristolochioides are reviewed. Ten different samples from Africa and Arabia, representing the main morphotypes, were analysed using RAPDs. The dendrogram resulting from cluster analysis is compared with the current taxonomy. Only two subspecies within C. aristolochioides should be recognized, despite considerable differences in floral morphology within both. It is proposed to confine the typical subspecies to tropical Africa and to consider C. aristolochioides ssp. albertina and Ceropegia seticorona as synonyms. C. aristolochioides ssp. deflersiana, easily distinguished by hairy carpels, is restricted to the SW Arabian Peninsula. The formal recognition of varieties within each subspecies, which might be considered with regard to the highly diverse flower morphology, is rejected.     

Sauropod dinosaurs from the Lower Cretaceous of eastern Asia: taxonomic and biogeographical implications

Sauropod dinosaurs are poorly represented in the Lower Cretaceous of eastern Asia. Here, we describe a number of isolated sauropod teeth from the Kuwajima Formation (?Berriasian–?Hauterivian) of Shiramine, Japan. The mosaic of shared derived characters and symplesiomorphies displayed by the teeth indicate that they are referable to a basal member of the titanosauriform radiation. A taxonomic review of previously described sauropod specimens from eastern and south–eastern Asia reveals that a diversity of sauropods (including a titanosaurian, a basal titanosauriform and a ?euhelopodid, as well as several forms of indeterminate systematic position) was present in this region in the Early Cretaceous. This diversity conflicts with previous suggestions that eastern Asia was biogeographically isolated from the rest of Laurasia until the late Early Cretaceous and that the sauropod fauna was limited to the endemic East Asian clade Euhelopodidae. The presence of titanosauriform sauropods in the basal Cretaceous of Japan and Thailand indicate that the proposed faunal isolation of eastern Asia ended approximately 20 myr earlier than usually suggested.     

Isolated theropod teeth from the Kem Kem Beds (Early Cenomanian) near Taouz, Morocco

Thirty-seven well-preserved, isolated theropod teeth from the Early Cenomanian Kem Kem beds, Morocco, are identified by using morphometric data and direct comparison with teeth previously described in the literature. Direct comparison reveals that four different morphotypes (MT 1–4) are present in the sample. The teeth of MT 1 are characterised by unserrated carinae and belong to spinosaurid dinosaurs. The teeth of MT 2–4 have serrated carinae, and our data analysis indicates they are of carcharodontosaurid, dromaeosaurid, and abelisaurid origin. Three types of crown enamel ornamentation are present among the teeth of MT 1, which implies that, apart from Spinosaurus aegyptiacus STROMER 1915, more than one species of spinosaurine theropods may be present in the Early Cenomanian of Northern Africa. Our results also confirm the occurrence of abelisaurids, dromaeosaurids, and carcharodontosaurids in Morocco.     

Tetrapod teeth from an Upper Ladinian bone bed, Schöningen (Lower Saxony, Germany)

A former clay quarry near Schöningen in Lower Saxony exposes deposits dating from the upper half of the Ladinian. There are several bone beds in this quarry, which differ partly in their faunal composition and in the preservation of the fossils. One of these bone beds contains many morphologically different teeth of mainly terrestrial tetrapods and a variety of remains of actinopterygians, chondrichthyans and dipnoans. The tetrapod teeth are described in this paper. A precise taxonomic determination is not possible, but the material appears to contain two temnospondyls, a synapsid and several species of archosaurs. Some tooth morphotypes can not be assigned, even at a high taxonomic level. The tooth assemblage, which is described in this paper, is briefly compared with published and unpublished data on teeth from the Middle and Upper Keuper of Central Europe.     

Relative numbers of selected bacterial forms in different regions of the cockroach hindgut

The relative numbers of fourteen microbial morphotypes in transmission electron micrographs of the hindgut of a cockroach, Eublaberus posticus, were counted and their distribution was analyzed statistically. The microbiota of three wall-associated regions (the anterior paunch, the posterior paunch, and the black band region) was clearly different from that of the gut lumen. The three wall fractions were also significantly different from each other. Only one of the fourteen types, prosthecate bacteria, appeared to be distributed randomly in the four fractions. The five main wall-associated morphotypes individually constituted up to 41% of the microbes in some micrographs. They included one type with the characteristic morphology of Methanospirillum. Six morphotypes rarely made up over 2% of the population, but were consistently present. The numbers of the remaining three morphotypes were quite variable between micrographs and between individual insects, but when present often made up 5–10% of the population.     

Pterosaur remains from the Cretaceous of Morocco

New pterosaur remains consisting of jaw fragments of toothless taxa and isolated teeth are described from the red beds of the Kern Kern region of southern Morocco. The stratigraphic position of those red beds is discussed and it is concluded that they are in all likelihood early Cenomanian in age. At least four taxa of pterodactyloid pterosaurs are present. The toothless jaw fragments are referred to the families ?Pteranodontidae, ?Azhdarchidae and Tapejaridae. Four different morphotypes can be distinguished among the isolated teeth. They are tentatively referred to the Ornithocheiridae. This assemblage reveals a high diversity of pterosaurs in Africa during the early Upper Cretaceous. The possible occurrence of tapejarids and anhanguerids indicates relationships with the somewhat older pterosaur assemblage from the Santana Formation (Aptian/Albian) of Brazil. If confirmed, the presence of azhdarchids and pteranodontids in the early Cenomanian suggests an early differentiation of these essentially late Late Cretaceous groups of large pterosaurs.