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Determining the relationships of mid-Late Cretaceous African taxa is central to understanding the timing and resultant palaeobiogeographical patterns of Gondwanan fragmentation. The early Late Cretaceous (Cenomanian) Kem Kem beds of southeastern Morocco preserve a diverse vertebrate fauna, including sauropod dinosaurs. Sauropod material includes the holotype of the rebbachisaurid diplodocoid Rebbachisaurus garasbae and fragmentary remains representing indeterminate titanosauriforms and rebbachisaurids. Here, we describe two new specimens from the Kem Kem beds. A dorsal neural arch with complex internal pneumaticity is tentatively attributed to a somphospondylan titanosauriform. A caudal vertebra possessing several rebbachisaurid synapomorphies is excavated by a large and pervasive lateral pneumatic foramen, a feature undocumented in other rebbachisaurids. However, caudal vertebrae are currently unknown for the sympatric R. garasbae, so this element could be referable to that taxon or a second, previously unknown, rebbachisaurid species. Interestingly, this new caudal vertebra displays a mosaic of features otherwise restricted to limaysaurine or nigersaurine rebbachisaurids, suggesting a placement basal to these clades, which is the position usually recovered for R. garasbae in phylogenetic analyses. A review of the mid-Cretaceous African sauropod fossil record removes the Cretaceous record of dicraeosaurids from Africa, restricting this clade to a single post-Jurassic occurrence in Argentina. All diagnostic sauropod remains can be attributed to titanosauriforms or rebbachisaurids. Whereas rebbachisaurids were seemingly restricted to northwestern Africa and disappeared post-Cenomanian, titanosauriforms were widespread across the African continent and survived until the latest Cretaceous. The development of the mid-Cretaceous Trans-Saharan Seaway might have acted as a dispersal barrier for rebbachisaurids and other vertebrate groups. In contrast, titanosauriforms might have been able to cross this barrier, but it is possible that they were also unable to disperse, and that northwestern African titanosauriforms were not closely related to taxa from the rest of the African continent. New materials and a better understanding of titanosaur interrelationships will be crucial in teasing these scenarios apart.  相似文献   

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The Early Cretaceous lizard Yabeinosaurus was one of the first taxa described from the now famous Jehol Biota of northeast China. Misinterpreted for more than 60 years and misclassified as a gekkotan based on juvenile specimens, it is now recognised to be a large, well-ossified lizard with an extended period of skeletal maturation. Here we describe two additional complete specimens of Yabeinosaurus that provide new information on skeletal morphology, most notably of the skull, pectoral girdle, and tail. Both specimens also preserve gut contents, showing that large individuals took vertebrate prey, including fish. A more complete understanding of Yabeinosaurus permits a review of the type specimen of Yabeinosaurus youngi. Skull traits used to distinguish Y. youngi from Yabeinosaurus tenuis are invalid but, as noted by Hoffstetter, the two species differ markedly in limb proportions. Attribution of Young’s specimen to Yabeinosaurus is equivocal, but could potentially extend the temporal range of the lineage into the Jurassic. A new phylogenetic analysis based on a morphological data set places Yabeinosaurus on the stem of Scleroglossa, as the sister taxon of the contemporaneous Japanese lizard Sakurasaurus.  相似文献   

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Yabeinosaurus was the first lizard genus described from the Early Cretaceous Jehol Biota of China. The holotype of the type species, Y. tenuis, is an immature skeleton but it has been lost for decades. A second species, Y. youngi, was erected based on another immature skeleton and is distinguished by its longer, more gracile limbs. In 2001, a juvenile skeleton from the Jingangshan locality of Liaoning was designated as the neotype for Y. tenuis. Subsequently, several further specimens from other localities, juvenile and adult, have been attributed to this species. Here we describe a second lizard specimen from the neotype locality. In anterior tooth shape, finger-like mandibular angular process, cranial sculpture, maxilla shape, and short, robust humerus, the new specimen resembles adult and subadult material currently attributed to Y. tenuis, but it differs in having bicuspid posterior teeth and a straight rather than a hooked angular process of the mandible. We propose a new species (Y. bicuspidens sp. nov.) for this specimen which represents the first occurrence of bicuspid teeth in an Early Cretaceous lizard from China. However, the recognition of a second species at the neotype locality raises a taxonomic problem. Due to the immaturity of the designated holotype and neotype of Y. tenuis, they cannot be coded for the jaw and dental characters that distinguish the two robust-limbed species. This renders Yabeinosaurus tenuis a nomen dubium. Here we propose that Yabeinosaurus tenuis should be treated as a historic taxon, permitting retention of the generic name under ICZN rules, and erect a second new species (Y. robustus sp. nov.) for the principal robust-limbed morphotype in the Jehol Biota with monocuspid posterior teeth and a hooked angular process.  相似文献   

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The Santa Elena peninsula in the northwest of Costa Rica protrudes about 30 km westwards into the Pacific Ocean, and measures about 8–16 km in a north–south direction. Several geological studies have been carried out since 1953, showing that much of the peninsula is made up of peridotite, cut by mafic dykes. Only one previous brief examination appears to have been made of the vegetation in relation to the composition of the soils. We present here the results of a survey of some soils and plants of the eastern part of the peridotite massif, in which 73 plant specimens representing 51 identified species were collected and analyzed. The soils sampled all showed extreme ultramafic characteristics: Fe 10–16%, Mg 4–16%, Ca 0.5–1.4%; Ni 3000–7500 mg/kg, Cr 1400–3650 mg/kg, Co 150–325 mg/kg. The plants collected include several from genera such as Arrabidaea, Chamaesyce, Helicteres, Hyptis, Lippia, Oxalis, Polygala, Turnera and Waltheria that are also represented on ultramafics elsewhere in the Americas (e.g. Cuba, Puerto Rico, Brazil). Few of the species appear to be endemic to Costa Rica or to the ultramafics of Santa Elena. None of the specimens collected exhibited hyperaccumulation of nickel, the highest Ni concentration being 275 mg/kg in Buchnera pusilla.  相似文献   

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Cu-Fe-sulphide mineral assemblages from the Olympic Dam (OD) Fe-oxide Cu-U-Au-Ag deposit, South Australia, are studied down to the nanoscale to explore the potential these minerals have for understanding genetic processes such as primary deposit zonation. Cu-Fe-sulphide pairs: ‘brown’ bornite associated with chalcopyrite (bornite-chalcopyrite zone); and symplectites of ‘purple’ bornite with species from the chalcocite group, Cu2  xS (bornite-chalcocite zone), co-define an upwards and inwards deposit-scale zonation at OD. In the bornite-chalcocite zone, there is also an increase in the proportion of chalcocite relative to bornite within the symplectites towards upper levels. In this case, two-phase Cu2  xS assemblages are also present, as anisotropic, hexagonal chalcocite (CcH) with lamellar exsolutions of digenite, distinguishable at the μm-scale. Using compositional data (electron microprobe) combined with Transmission Electron Microscopy (TEM) study of foils prepared in–situ via Focused Ion Beam (FIB)-SEM, we show that Cu-Fe-sulphides from different ore zones feature nanoscale intergrowths, lattice defects, superstructure domains (na) and antiphase boundary domains (APBs) that can be interpreted as due to exsolution, coarsening and phase transformation during cooling from high-T solid solutions in the system Cu-Fe-S and sub-systems according to published phase diagrams. ‘Brown’ bornite [(Cu + Fe)/S > 5] contains pervasive lamellae of chalcopyrite which extend down to the nanoscale; such specimens appear homogeneous at the μm-scale. ‘Purple bornite’ [(Cu + Fe)/S < 5] in high-bornite symplectites is associated with chalcocite that shows APBs with 6a digenite and low-T chalcocite. Comparable APBs are also found in the ‘chalcocite’ zone with apparent homogeneity at the μm-scale. Both bornites contain exsolutions of djurleite. Systematic variation of Me/S and Cu/Fe in the two types of bornite points, however, to distinct origins from different bornite solid-solutions in the system Cu-Fe-S. Both show 2a and 4a intermediate superstructures. High-order superstructures (6a and incommensurate na) are restricted to the ‘purple’ bornite whereas the 2a4a low-T superstructure is found in both cases. Me/S ratios in the chalcocite group are variable; lower ratios (down to 1.8; digenite) are more common in chalcocite from symplectites with ‘purple’ bornite. Me/S can be as low as 1.4 where associated with ‘blue’ varieties (‘blaubleibender covellin’) of replacement origin. The two-phase Cu2  xS associations contain hexagonal chalcocite (Me/S = 1.95), lamellae of Cu-rich digenite (Me/S = 1.92), and anilite (Cu7S4) as nm-scale lamellae. Digenite shows 3a and 6a superstructures and CcH shows transition to pseudo-orthorhombic chalcocite. The presence of superstructures, high-T species and APBs is evidence for Cu-(Fe)-sulphide formation from high-T solid solutions at T > 300 °C (high-T phases, Cu-poor digenite), followed by cooling along distinct paths down to < 120 °C (APBs). The scenario of ‘exsolution from primary solid-solution’, corroborated by the consistency in phase relations within each zone across different scales of observation from deposit scale to nanoscale, backs up a model of primary hypogene ore precipitation rather than replacement, and accounts for the observed vertical zoning at OD. The FIB-TEM approach here is readily applicable to other deposits and shows that nanoscale observations are a valuable, although often overlooked, source of information to constrain ore genesis.  相似文献   

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Two new genera from the Lower Cretaceous Yixian Formation of Beipiao City, Liaoning Province, northeastern China are described and illustrated. Paracretocateres gen. nov. (type species P. bellus, sp. nov.) and Yixianteres gen. nov. (type species Y. beipiaoensis sp. nov.) are assigned to the subfamily Lophocaterinae based on the state of the procoxae which are not projecting, the transverse procoxal cavities, and the exposed protochantins. The new taxa broaden the diversity of this family and provide further morphological characters for phylogenetic studies of Trogossitidae. They also provide evidence that lophocaterines well-diversified prior to the trogossitines and their biologies may have been similar to extant forms.  相似文献   

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Univariate and multivariate statistics were applied to analyse the morphometrical variability of 4920 upper cheek teeth (P4, M1 and M2) of cave bears from 123 geographical sites (180 samples) of different Pliocene – Pleistocene ages. The analysed specimens included those belonging to the big cave bears Ursus kudarensis, Udeningeri, Uspelaeus (three subspecies) and Ukanivetz (including Uingressus), as well as the small cave bear Urossicus. The information‐theoretical parameters (Shannon entropy and orderliness (Von Foerster, 1960: On self‐organizing systems and their environments. In Self‐Organizing Systems, 31–50. Pergamon Press, London) were used to estimate tooth diversity in different teeth, different taxa and in selected local chrono‐populations. Multivariate allometry coefficients (Klingenberg, 1996: Multivariate allometry. In Advances in Morphometrics, 23‐49. Plenum Press, New York) were used to describe the relationships of different ‘parts’ of a tooth and to compare allometric patterns amongst species or selected local samples. A multivariate analysis showed a significant overlap of the size/shape parameter ranges in deningeroid and spelaeoid bears within morphological spaces. Within the cave bear lineage, the Deninger's bear has the greatest morphological diversity index (entropy) of all the teeth overall, and the lowest diversity is observed in the final taxon of this lineage – Ukanivetz (=ingressus). The P4 and M2 diversity showed multidirectional correlations with elevation above sea level amongst several ‘local’ populations of Late Pleistocene cave bears. The morphological disparities between the studied taxa are in close agreement with the distances in the available schemes of genetic differentiation based on ancient mitochondrial DNA. The split of Ukudarensis and Udeningeri has a good bootstrap support, which corresponds to the hypothesis about their parallel evolution. The small cave bear Urossicus is placed between Uarctos and Udeningeri. The phylogenetic signal is more pronounced in the variability of teeth in comparison with other skeletal remains of cave bears (cranium, mandible, or metapodial bones).  相似文献   

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Two fossils belonging to a new genus and species of water measurer (Gerromorpha, Hydrometridae), Alavametra popovi Sánchez-García and Nel gen. n., sp. n., are described as first definitive record of the family in Lower Cretaceous (upper Albian) amber from the Utrillas Group (Peñacerrada I site) in Spain. Although several parts of the specimens are obscured due to preservation, a sufficient number of taxonomical characters are visible to consider adequate placement within Heterocleptinae, including the very long posterior pair of cephalic trichobothria inserted on tubercles and the preapical articulation between the first and second antennal segments. The new fossil taxon is included into a cladistic analysis with extinct and extant hydrometrids, and it results putatively basal among the subfamily Heterocleptinae, suggesting that this clade was already present 105 Ma ago.  相似文献   

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《Cretaceous Research》2008,29(1):115-130
The diminutive (2.5–3.0 cm long), Cretaceous dinosaur track ichnogenus Minisauripus, previously known only from the type ichnospecies, M. chuanzhuensis, from a single locality in Sichuan Province China, is here reported from two new localities in South Korea and one in China. Material from the new Chinese locality is assigned to the new ichnospecies M. zhenshuonani on the basis of its distinctive morphology. Most of the new material is well-preserved, revealing narrow asymmetric tracks with claw traces, long step and phalangeal formula (2-3-4 for digits II, III and IV, respectively), suggesting a theropod track maker rather than an ornithischian, as originally inferred for the Chinese type material.The South Korean samples (eight tracks), from two localities in the Haman Formation, are considered Early Cretaceous (Aptian-Albian) in age, whereas the Chinese type material (21 specimens) has been assigned both an Early and Late Cretaceous age. The former age is probably correct as suggested by a new Minisauripus locality (5 specimens) from the Early Cretaceous (Barremian-Albian) of Shandong Province, China.Other diminutive tracks from the Sichuan fauna include Aquatilavipes sinensis (2.5 cm long, a possible junior synonym of Koreanoris hamanensis), Grallator emeiensis (2 cm long) and Velociraptorichnus sichuanensis (10–11 cm), which occur, in various combinations, with Minisauripus at both the new Korean and Chinese localities.In Minisauripus, digit III is very short in comparison with other theropods and provides a striking contrast to G. emeinsis. This difference has significant implications for standard assumptions about theropod track allometry. Based on the classic Early Jurassic forms Grallator and Eubrontes, it has long been inferred that relative digit III length shrank with increasing size (up to foot lengths of 30–40 cm). The reiteration of reduction in relative length of digit III in specimens in the size range of 2–3 cm indicates that the allometric or morphodynamic ‘program’ that influenced development in large theropod clades reiterated fractally in theropod clades a full order of magnitude smaller. This shows that a given allometry can be size-dependent in one clade and size-independent in another. Thus, the developmental program appears ‘contracted’ or morphologically miniaturized by heterochrony to manifest paedomorphically in some clades and peramorphically in others. This strongly suggests that ‘formal’ developmental ‘programs operated’ along similar morphodynamic lines in quite different clades.  相似文献   

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Ice sheets flowing across a sedimentary bed usually produce a landscape of blister-like landforms streamlined in the direction of the ice flow and with each bump of the order of 102 to 103 m in length and 101 m in relief. Such landforms, known as drumlins, have mystified investigators for over a hundred years. A satisfactory explanation for their formation, and thus an appreciation of their glaciological significance, has remained elusive. A recent advance has been in numerical modelling of the land-forming process. In anticipation of future modelling endeavours, this paper is motivated by the requirement for robust data on drumlin size and shape for model testing.From a systematic programme of drumlin mapping from digital elevation models and satellite images of Britain and Ireland, we used a geographic information system to compile a range of statistics on length L, width W, and elongation ratio E (where E = L/W) for a large sample. Mean L, is found to be 629 m (n = 58,983), mean W is 209 m and mean E is 2.9 (n = 37,043). Most drumlins are between 250 and 1000 metres in length; between 120 and 300 metres in width; and between 1.7 and 4.1 times as long as they are wide. Analysis of such data and plots of drumlin width against length reveals some new insights. All frequency distributions are unimodal from which we infer that the geomorphological label of ‘drumlin’ is fair in that this is a true single population of landforms, rather than an amalgam of different landform types. Drumlin size shows a clear minimum bound of around 100 m (horizontal). Maybe drumlins are generated at many scales and this is the minimum, or this value may be an indication of the fundamental scale of bump generation (‘proto-drumlins’) prior to them growing and elongating. A relationship between drumlin width and length is found (with r2 = 0.48) and that is approximately W = 7 L 1/2 when measured in metres. A surprising and sharply-defined line bounds the data cloud plotted in EWL space, and records a scale-dependent maximum elongation limit (approximated by Emax = L1/3, when L measured in metres). For a given length, for some reason as yet unknown, drumlins do not exceed the elongation ratio defined by this scaling law. We also report and compare our statistics to an amalgamated sample (25,907 drumlins) of measures derived from around 50 published investigations. Any theory must be able to explain the drumlin statistics and fundamental scaling properties reported herein and they thus provide powerful tests for drumlin modelling.  相似文献   

14.
The holotype of ‘Procheneosaurusconvincens, a juvenile lambeosaurine, is the most complete hadrosaurid known from Kazakhstan. North American species of Procheneosaurus are considered juveniles of Corythosaurus, Hypacrosaurus, and Lambeosaurus, rendering the generic name unusable. A replacement name, Kazaklambia convincens comb. nov., is herein proposed as this specimen is morphologically distinct from other Eurasian taxa and known juvenile lambeosaurines at a similar ontogenetic stage in having a prefrontal process of the postorbital with a dorsal thickening forming a dome lateral to the frontal dome, doming of the nasal anterodorsal to the orbit, and a frontal length/width ratio <1. The juvenile status of Kazaklambia makes phylogenetic placement difficult; however, morphometric and morphological information (particularly in relation to the hollow cranial crest and the length of the frontal) suggest a close affiliation with the basal lambeosaurines Amurosaurus and Tsintaosaurus, and support the hypothesis for an Asian origin for Lambeosaurinae.  相似文献   

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There has been considerable debate about whether the controversial tyrannosauroid dinosaur ‘Nanotyrannus lancensis’ from the uppermost Cretaceous of North America is a valid taxon or a juvenile of the contemporaneous Tyrannosaurus rex. In a recent Cretaceous Research article, Schmerge and Rothschild (2016) brought a new piece of evidence to this discussion: the morphology of the dentary groove, a depression on the lateral surface of the dentary that houses neurovascular foramina. They argued that an alleged ‘Nanotyrannus’ specimen, which possesses a groove, cannot be referable to Tyrannosaurus rex, which they considered as lacking the groove, and they hypothesized that ‘Nanotyrannus’ is closely related to albertosaurine tyrannosauroids, which also are said to possess the groove. However, we show that the groove is a widespread feature of tyrannosauroids that is present in T. rex and many other specimens, and that it is an ontogenetically variable feature that changes from a sharp, deeply-impressed groove to a shallower sulcus as an individual matures. As a result, the presence or absence of a dentary groove does not clarify the validity of ‘Nanotyrannus’ or its phylogenetic position among tyrannosauroids. We consider it most parsimonious that ‘Nanotyrannus’ specimens belong to juvenile T. rex.  相似文献   

16.
The cymatoceratid nautilid genus Anglonautilus is distinguished from most other post-Triassic nautiloids by the occurrence of pronounced fold-like undulating ribs on the phragmocone and early body chamber. Anglonautilus praeundulatus n. sp. is described from the lower Aptian of eastern Spain. It is the first record of this genus from Spain and constitutes the oldest definite representative of the genus. This places the new species at the base of the evolutionary history of Anglonautilus. Its ornamental features confirm the previously assumed close relationship between Anglonautilus and Cymatoceras. An analysis of the ornamental pattern of all species hitherto referred to Anglonautilus indicates that there is an evolutionary lineage leading to the type species of the genus, A. undulatus, and a single successor (A. subalbensis). Several Late Cretaceous species hitherto identified as Anglonautilus (A. japonicus, A. mamiyai, A. suciensis) bear fold-like ribs superficially similar to typical representatives of Anglonautilus. The pattern of ornament in these taxa is very different at closer inspection, though. This suggests that these species developed independently from Cymatoceras. The undulations present in these taxa are interpreted here as a result of convergent evolution. They are therefore excluded from the genus Anglonautilus and provisionally referred to as “Anglonautilus” spp.  相似文献   

17.
The first alveolinoidean appearing in the Cenomanian Natih Formation of Oman (Adam foothills and southern edge of Jabal Akhdar) are studied in detail. Morphological analysis results in the creation of one new family, Myriastylidae, two new genera, Myriastyla and Alveocella, and four new species, M. omanensis, M. grelaudae, A. wernliana, and Cisalveolina nakharensis. These four new taxa have a short stratigraphic extension restricted to the uppermost part of Natih E unit and are dated early middle Cenomanian by neighboring foraminifers and ammonite levels.  相似文献   

18.
Cenomanian rocks of Egypt are rich in moderately to well-preserved echinoids. From the large Cretaceous echinoid collection at the Cairo Geological Museum, a total of 27 regular species, in 12 genera, eight families, six orders and two subclasses, are revised and systematically described from these strata as exposed at several localities in Egypt. One new family (Pedinopsidae), one new subfamily (Lorioliinae), one new genus (Alternocidaris) and two new species of Goniopygus, G. subaequalis (Gebel Ekma) and G. macrotuberculatus (Gebel El Minsherah), are erected. Additionally, two species of Diplopodia, D. micropyga Fourtau, 1921 and D. halperti Fourtau, 1921, are transferred to the genus Pedinopsis.  相似文献   

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The taxonomic status of specimens referred to Tarchia kielanae, T. gigantea, and Minotaurasaurus ramachandrani has been contested. The holotype of T. kielanae, Zaklad Paleobiologii (Institute of Paleobiology)-Polish Academy of Sciences (ZPAL) MgD I/111, is redescribed. It has features common to both Palaeontological Institute-Russian Academy of Sciences (PIN) 3142/250, long considered Tarchia, and to M. ramachandrani but which are lacking in Saichania chulsanensis. The specimen PIN 3142/250 is not referable to Saichania but instead represents a new species, Tarchia teresae sp. nov. Similarly, the holotype skull of M. ramachandrani is not referable to T. kielanae, so Minotaurasaurus is a valid taxon. Tarchia is more derived than either Saichania or Minotaurasaurus while sharing features of both.  相似文献   

20.
A new genus and species of small (3.5 mm excluding ovipositor) parisitoid wasp is described from the Lower Cretaceous (Aptian) Crato Formation Lagerstätte of Brazil. Parviformosus wohlrabeae gen. et sp. nov. is known from a single female imago and is assigned to Pteromalidae. It is diagnosed by the robustness of the scutellum, the structure, size and positioning of the mesopleuron, the complexity of the propodeum–petiole junction and a posteriorly curved dorsal ‘lip’ on metasomal segment 4. At only 3.5 mm in length, P. wohlrabeae is the smallest fossil wasp from the Cretaceous of South America and the first Mesozoic representative of Pteromalidae.  相似文献   

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