Distribution of the dentary groove of theropod dinosaurs: Implications for theropod phylogeny and the validity of the genus Nanotyrannus Bakker et al., 1988

This study examines the phylogenetic distribution of a morphologic character, described as a groove containing pores, on the lateral surface of the dentary bone in theropod dinosaurs. The nature of this groove is a feature unique to theropods. Of the 92 theropod taxa examined for the presence and absence of this feature, 48 possessed and 44 lacked this feature. Distribution of this character was compared to published phylogenetic analyses of theropods, in order to evaluate the utility of the dentary groove as a diagnostic feature. 80% of pre-Tyrannoraptoran theropods possessed the dentary groove, with only 6 reversals in basal theropod clades. Theropods with beaks or edentulous jaws all lacked a dentary groove. Tyrannosauroidea is marked by mosaic distribution of this character. Among tyrannosauroids, the dentary groove occurs only in Dryptosaurus and the Albertosaurinae (Albertosaurus + Gorgosaurus). Nanotyrannus lancensis, sometimes described as representing juvenile Tyrannosaurus rex, also possesses this groove, unlike the remainder of the Tyrannosaurinae. Nanotyrannus lancensis was included in a phylogenetic analysis of Tyrannosauroidea and was recovered within Albertosaurinae. We recommend that Nanotyrannus stand as a valid taxon nested within the Albertosaurinae, based on the presence of this groove, as well as other features of the skull.     

A juvenile Edmontosaurus from the late Maastrichtian (Cretaceous) of North America: Implications for ontogeny and phylogenetic inference in saurolophine dinosaurs

The anatomy of an articulated juvenile specimen of the saurolophine hadrosaurid dinosaur Edmontosaurus annectens, LACM 23504, is described in detail. This individual consists of a partial skull and nearly complete articulated postcranium, collected from upper Maastrichtian strata of the Hell Creek Formation of Montana, USA. This study aims to help understand the ontogenetic morphological changes occurring in the cranium and postcranium of saurolophine hadrosaurids using E. annectens as a case study. Emphasis is placed on those morphological attributes that are ontogenetically variable via comparative osteology between the juvenile and the available adult specimens. It is observed that much of the cranial ontogenetic variation relates to the elongation of the skull and mandible. In the postcranium, most of the ontogenetic variation concentrates in the pectoral and pelvic girdles and the stylopodia. Not all the identified patterns of ontogenetic variation may be generalized to all hadrosaurids. The impact of ontogenetic variation on phylogenetically informative characters of saurolophine hadrosaurids is evaluated. It is concluded that, at least for Edmontosaurus annectens (and perhaps other saurolophine hadrosaurids), most characters used in phylogenetic inference of these animals are not affected by ontogeny. Thus, juvenile specimens are still a source of substantial character data suitable for use in phylogenetic analyses of saurolophine relationships. Nevertheless, it is recommended that ontogenetically variable characters are left as missing data in a character–taxon matrix when only juvenile material is available for a given saurolophine taxon. Scoring those characters based solely on juveniles is likely to decrease the accuracy of the phylogenetic inference.     

Description of a juvenile titanosaurian dinosaur from the Upper Cretaceous of Brazil

We describe a juvenile specimen of a titanosaurian sauropod dinosaur consisting of two dorsal and three caudal vertebral centra, an ilium fragment, and an ischium unearthed in 1991 from Site Km 153.5 at BR-050 highway in the Serra da Galga region, municipality of Uberaba, state of Minas Gerais, Brazil. The outcrop at the site is assigned to the Serra da Galga Member of the Marília Formation (Bauru Basin: Bauru Group; Upper Cretaceous: Maastrichtian). Although the material is very incomplete, features such as strongly procoelous caudal centra suggest an affinity with the titanosaurian clade Lithostrotia. The extensive vertebral pneumaticity with deep pleurocoels and well-developed camerae supports the hypothesis that, in titanosaurs, the air sac system was already present and fully developed even at early ontogenetic stages.     

Long bone histology of a eusuchian crocodyliform from the Upper Cretaceous of Spain: Implications for growth strategy in extinct crocodiles

The long bone histology of a Late Cretaceous eusuchian crocodyliform from the Iberian Peninsula reveals clear variations in the cortical structure which reflects changes in the speed of bone deposition (i.e., skeletal growth) related to ontogeny. The presence of secondary woven-fibred bone tissue in the perimedullar region of the cortex, and the existence of an external fundamental system in the most external periostic cortex, which is a proxy for somatic maturity and effective cessation of growth, challenges the former idea that the growth strategy of extinct crocodylians fit in the typical ectotherm condition, according to which these animals grew slowly during life under an indeterminate growth strategy. The analysed specimen lived for a minimum of 16 years and the highest preserved apposition rates took place in an advanced ontogenetic stage. The study suggests that the general aspects of the modern crocodylian growth strategy were already in place in some lineages by the Cretaceous.     

Re-characterization of Tylosaurus nepaeolicus (Cope, 1874) and Tylosaurus kansasensis Everhart, 2005: Ontogeny or sympatry?

Tylosaurus nepaeolicus (Cope, 1874), from the lower Smokey Hill Chalk upper Coniacian of Kansas is reassessed and compared to T. kansasensis Everhart, 2005, its sympatric contemporary from the same formation; both are compared to a later species from the upper Smoky Hill Chalk, T. proriger (Cope, 1869). Tylosaurus nepaeolicus (Cope, 1874) is virtually indistinguishable from T. kansasensis Everhart, 2005, and both show important similarities with T. proriger, particularly the smaller individuals of T. kansasensis. Many of the anatomical features of T. kansasensis are indicative of a juvenile stage based on comparisons to T. proriger. In addition to the aforementioned spatial and temporal sympatry between T. nepaeolicus and T. kansasensis, it is anatomically difficult to distinguish the two species from each other, with the few notable differences being ontogenetically variable, and possibly indicating allometric changes during post-embryonic development; in addition, T. nepaeolicus is known from fragmentary remains of very large individuals, while T. kansasensis is known from a small number of complete and recently collected skulls, though of much smaller size than the type materials of T. nepaeolicus. We suggest that T. kansasensis is a juvenile of Tylosaurus nepaeolicus, and is thus the junior synonym of the latter. Furthermore, we find ontogenetic evidence to suggest that T. proriger is likely a paedomorph of T. nepaeolicus, albeit, a gigantic one.     

Dentary groove morphology does not distinguish ‘Nanotyrannus’ as a valid taxon of tyrannosauroid dinosaur. Comment on: “Distribution of the dentary groove of theropod dinosaurs: Implications for theropod phylogeny and the validity of the genus Nanotyrannus Bakker et al., 1988”

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.     

Dimorphism of Bivalved Arthropod Branchiocaris? Yunnanensis from the Early Cambrian Chengjiang Biota, South China

The bivalved arthropod Branchiocaris? yunnanensis Hou, 1987 is redescribed on the basis of new fossil material from the early Cambrian Chengjiang biota, South China. In total, 81 specimens have been examined. The carapace comprises two identical valves, each valve are sub-circular in outline, ranging from 24 mm to 58 mm in length, and from 15 mm to 46 mm in height. The dorsal margin is straight and bears two short cardinal spines. The valve surface is either smooth or ornamented with polygonal reticular structures, which may result from different preservation conditions. Two morphotypes have been recognized based on the presence or absence of a subdorsal swelling of the valve, which is convex dorsolaterally and extends beyond the dorsal margin when it appears. Statistic and ontogenetic analyses show both morphotypes grow isometrically and have the same growth trajectory. Therefore, the two morphotypes are interpreted as dimorphism within the same species. Our results are helpful for investigating the taxonomy and ontogeny of arthropod fossils and suggest that dimorphism might be fairly common in these early arthropods.     

泥蚶(Tegillarca granosa)个体发育过程中同工酶基因表达与调控的研究

采用不连续垂直聚丙烯酰胺凝胶电泳法，对泥蚶(Tegillarca granosa Linnaeus)19个发育阶段的ADH、LDH、EST、IDH、ALP、MDH、GDH、POD、ATPase和ME等10种同工酶的基因表达进行检测分析。结果表明，泥蚶在个体发育过程中随着机体分化和生理活动的改变，同工酶基因的表达呈现出显著的时序变化。在10种酶中，LDH和ATPase两种酶的所有酶带及EST、MDH和ME等的部分酶带在各个发育阶段持续表达，其他酶带只出现在发育阶段的某一或某些阶段，并且表现出了与泥蚶营养方式、代谢形式等生理活动的关联性。在泥蚶的个体发育阶段过程中，IDH、MDH、GDH、ME、ALP等5种酶在32细胞期表达的酶带数剧增，因此32细胞期可能是合子基因表达启动的时间；ADH、EST、LDH、MDH、GDH、ME等6种酶在泥蚶的附着变态期酶带大幅减少或产生新酶带，可能与泥蚶附着期运动方式、代谢强度及生活环境的改变有关。     

Osteology and ontogeny of Early Cretaceous Philydrosaurus (Diapsida: Choristodera) based on new specimens from Liaoning Province,China

Choristoderes are a group of extinct diapsid reptiles that once occupied the freshwater systems in the Northern Hemisphere from the Middle Jurassic through the Miocene. The Early Cretaceous monjurosuchid Philydrosaurus from western Liaoning, China, represents a transitional morphotype between a broad-snouted (crocodile-like) and a narrow-snouted (gavial-like) skull during the evolution of Choristodera. New specimens of the taxon from the Early Cretaceous Jiufotang Formation include an incomplete braincase and a nearly complete juvenile skeleton; the study of these specimens provides new information on the braincase and ontogenetic change of the skeleton of this transitional monjurosuchid. In the braincase floor, the foramen internus canalis caroticus externus opens in a groove lateral to the constricted lateral aspect of the parasphenoid, and ventral exposure of this foramen differs from Champsosaurus, in which parasphenoid/pterygoid fusion completely encloses the canal. On the occiput, the vagus nerve foramen penetrates the basioccipital, while the two foramina for the hypoglossal nerve open at the exoccipital/basioccipital suture. Comparative study of the new juvenile specimen with adult Philydrosaurus reveals previously unknown developmental changes of the cranial and postcranial skeleton of this monjurosuchid. Ontogenetically, Philydrosaurus underwent a dramatic change of skull proportions, including elongation of the antorbital and postorbital regions, and elongation of the jaws, with a great increase of the number of marginal teeth. The lower temporal fenestra is entirely closed in early ontogeny, as in large, fully-grown adults.     

Autotrophic response to lake age,conductivity and temperature in two West Greenland lakes

Predicted changes in future climate necessitate a better understanding of climate impacts on lake biota, and the role of within-lake processes in modifying biotic response. Therefore we examined two climate-related variables (lake-water conductivity and GRIP temperatures) and lake ontogeny (lake age), to determine their influence on lake autotrophic communities in two neighbouring closed-basin lakes from West Greenland spanning the past 8,000 years. Using sedimentary pigments as proxies for lake autotrophic communities, we used synchrony and variance partitioning analyses (VPA) to test three specific hypotheses (a) that lake primary production would increase with lake age, (b) that climate would be the dominant process controlling autotrophic communities in these pristine lakes and (c) that the response of autotrophs to conductivity and temperature would vary depending upon the age of the lake. The results supported our first hypothesis, showing that lakes changed significantly with age, exhibiting an increase and decline in production in the first millennium of their existence, followed by a steady increase in production and increasingly frequent abrupt switches between mixed and meromictic states. The highly synchronous detrended response (r = 0.769) of lake autotrophs in the two study lakes, supported our hypothesis that climate was the dominant factor controlling lake autotrophs. However, VPA revealed that our climate-related variables (temperature and conductivity) explained only small amounts of variance alone (<12.9%) because covariance among them hindered efficient partitioning. In support of our third hypothesis, autotrophs responded significantly to temperature and conductivity in interaction with lake age (>50% variance explained) and with each other (>28% variance explained), such that autotrophic response changed as lakes aged. In spite of this, lakes sometimes responded independently, as a result of differences in the relative proportion of benthic to pelagic production and because of differences in lake morphometry. Together these results show that long-term control of lake autotrophs by climate and lake age is modified on shorter timescales by non-linear responses related to within-lake processes, and by the interaction of different climate variables with each other and with lake age.