The origin and early evolution of dinosaurs

The oldest unequivocal records of Dinosauria were unearthed from Late Triassic rocks (approximately 230 Ma) accumulated over extensional rift basins in southwestern Pangea. The better known of these are Herrerasaurus ischigualastensis, Pisanosaurus mertii, Eoraptor lunensis, and Panphagia protos from the Ischigualasto Formation, Argentina, and Staurikosaurus pricei and Saturnalia tupiniquim from the Santa Maria Formation, Brazil. No uncontroversial dinosaur body fossils are known from older strata, but the Middle Triassic origin of the lineage may be inferred from both the footprint record and its sister‐group relation to Ladinian basal dinosauromorphs. These include the typical Marasuchus lilloensis, more basal forms such as Lagerpeton and Dromomeron, as well as silesaurids: a possibly monophyletic group composed of Mid‐Late Triassic forms that may represent immediate sister taxa to dinosaurs. The first phylogenetic definition to fit the current understanding of Dinosauria as a node‐based taxon solely composed of mutually exclusive Saurischia and Ornithischia was given as “all descendants of the most recent common ancestor of birds and Triceratops”. Recent cladistic analyses of early dinosaurs agree that Pisanosaurus mertii is a basal ornithischian; that Herrerasaurus ischigualastensis and Staurikosaurus pricei belong in a monophyletic Herrerasauridae; that herrerasaurids, Eoraptor lunensis, and Guaibasaurus candelariensis are saurischians; that Saurischia includes two main groups, Sauropodomorpha and Theropoda; and that Saturnalia tupiniquim is a basal member of the sauropodomorph lineage. On the contrary, several aspects of basal dinosaur phylogeny remain controversial, including the position of herrerasaurids, E. lunensis, and G. candelariensis as basal theropods or basal saurischians, and the affinity and/or validity of more fragmentary taxa such as Agnosphitys cromhallensis, Alwalkeria maleriensis, Chindesaurus bryansmalli, Saltopus elginensis, and Spondylosoma absconditum. The identification of dinosaur apomorphies is jeopardized by the incompleteness of skeletal remains attributed to most basal dinosauromorphs, the skulls and forelimbs of which are particularly poorly known. Nonetheless, Dinosauria can be diagnosed by a suite of derived traits, most of which are related to the anatomy of the pelvic girdle and limb. Some of these are connected to the acquisition of a fully erect bipedal gait, which has been traditionally suggested to represent a key adaptation that allowed, or even promoted, dinosaur radiation during Late Triassic times. Yet, contrary to the classical “competitive” models, dinosaurs did not gradually replace other terrestrial tetrapods over the Late Triassic. In fact, the radiation of the group comprises at least three landmark moments, separated by controversial (Carnian‐Norian, Triassic‐Jurassic) extinction events. These are mainly characterized by early diversification in Carnian times, a Norian increase in diversity and (especially) abundance, and the occupation of new niches from the Early Jurassic onwards. Dinosaurs arose from fully bipedal ancestors, the diet of which may have been carnivorous or omnivorous. Whereas the oldest dinosaurs were geographically restricted to south Pangea, including rare ornithischians and more abundant basal members of the saurischian lineage, the group achieved a nearly global distribution by the latest Triassic, especially with the radiation of saurischian groups such as “prosauropods” and coelophysoids.     

Using creation science to demonstrate evolution: application of a creationist method for visualizing gaps in the fossil record to a phylogenetic study of coelurosaurian dinosaurs

It is important to demonstrate evolutionary principles in such a way that they cannot be countered by creation science. One such way is to use creation science itself to demonstrate evolutionary principles. Some creation scientists use classic multidimensional scaling (CMDS) to quantify and visualize morphological gaps or continuity between taxa, accepting gaps as evidence of independent creation and accepting continuity as evidence of genetic relatedness. Here, I apply CMDS to a phylogenetic analysis of coelurosaurian dinosaurs and show that it reveals morphological continuity between Archaeopteryx, other early birds, and a wide range of nonavian coelurosaurs. Creation scientists who use CMDS must therefore accept that these animals are genetically related. Other uses of CMDS for evolutionary biologists include the identification of taxa with much missing evolutionary history and the tracing of the progressive filling of morphological gaps in the fossil record through successive years of discovery.     

A new species of Azendohsaurus (Diapsida: Archosauromorpha) from the Triassic Isalo Group of southwestern Madagascar: cranium and mandible

Abstract: Here, we describe a new species of Azendohsaurus from the Middle–Late Triassic of Madagascar, extending the geographical range of a taxon known otherwise only by a single species from Morocco. Although Azendohsaurus has consistently been regarded as an early dinosaur (based on various advanced dental and gnathic features resembling those characterizing certain dinosaur subgroups), the relatively complete skeletal material, now available from Madagascar, argues strongly against its dinosaurian affinities. Rather, the retention of numerous primitive cranial and postcranial features indicates a surprisingly early divergence of Azendohsaurus within Archosauromorpha and an unusual mosaic of characters in this taxon. Features considered diagnostic of Sauropodomorpha thus are inferred to occur homoplastically in at least one clade of nondinosaurian archosauromorphs, indicating a complex evolution and distribution of features traditionally thought to be derived within archosaurs. Azendohsaurus has teeth resembling those of both early sauropodomorph and ornithischian dinosaurs, yet also possesses numerous inarguable basal archosauromorph cranial and postcranial attributes. This highlights the risk of uncritically referring isolated, Middle–Late Triassic (or even later), ‘leaf‐shaped’ teeth with denticles to the Dinosauria. Similarly, the occurrence of such teeth in an early diverging archosauromorph indicates that specializations for herbivory originated more frequently within this clade than conventionally assumed. For example, Azendohsaurus and numerous basal sauropodomorph dinosaur taxa share an array of convergently acquired features associated with herbivory, including tooth denticles, expanded tooth crowns, a downturned dentary and the articular located at the ventral margin of the mandible. Some of these features (denticles, expanded crowns and the ventrally deflected articular) are even more widespread among archosauromorphs, including aetosaurs, silesaurs and ornithischian dinosaurs. A downturned dentary also occurs in Trilophosaurus, a taxon further marked by unique specializations for herbivory, including transversely lophate, tricuspid teeth. An array of features associated with herbivory also occurs in rhynchosaurs and certain crocodilians (e.g. Simosuchus). This distribution suggests that craniodental features associated with herbivory were much more pervasive across the archosauromorph clade than previously recognized, possibly evolving at least six to eight times independently.     

Lower limits of ornithischian dinosaur body size inferred from a new Upper Jurassic heterodontosaurid from North America

The extremes of dinosaur body size have long fascinated scientists. The smallest (<1 m length) known dinosaurs are carnivorous saurischian theropods, and similarly diminutive herbivorous or omnivorous ornithischians (the other major group of dinosaurs) are unknown. We report a new ornithischian dinosaur, Fruitadens haagarorum, from the Late Jurassic of western North America that rivals the smallest theropods in size. The largest specimens of Fruitadens represent young adults in their fifth year of development and are estimated at just 65–75 cm in total body length and 0.5–0.75 kg body mass. They are thus the smallest known ornithischians. Fruitadens is a late-surviving member of the basal dinosaur clade Heterodontosauridae, and is the first member of this clade to be described from North America. The craniodental anatomy and diminutive body size of Fruitadens suggest that this taxon was an ecological generalist with an omnivorous diet, thus providing new insights into morphological and palaeoecological diversity within Dinosauria. Late-surviving (Late Jurassic and Early Cretaceous) heterodontosaurids are smaller and less ecologically specialized than Early (Late Triassic and Early Jurassic) heterodontosaurids, and this ecological generalization may account in part for the remarkable 100-million-year-long longevity of the clade.     

The oldest dinosaur? A Middle Triassic dinosauriform from Tanzania

The rise of dinosaurs was a major event in vertebrate history, but the timing of the origin and early diversification of the group remain poorly constrained. Here, we describe Nyasasaurus parringtoni gen. et sp. nov., which is identified as either the earliest known member of, or the sister–taxon to, Dinosauria. Nyasasaurus possesses a unique combination of dinosaur character states and an elevated growth rate similar to that of definitive early dinosaurs. It demonstrates that the initial dinosaur radiation occurred over a longer timescale than previously thought (possibly 15 Myr earlier), and that dinosaurs and their immediate relatives are better understood as part of a larger Middle Triassic archosauriform radiation. The African provenance of Nyasasaurus supports a southern Pangaean origin for Dinosauria.     

Fusion Patterns in the Skulls of Modern Archosaurs Reveal That Sutures Are Ambiguous Maturity Indicators for the Dinosauria

The sutures of the skulls of vertebrates are generally open early in life and slowly close as maturity is attained. The assumption that all vertebrates follow this pattern of progressive sutural closure has been used to assess maturity in the fossil remains of non-avian dinosaurs. Here, we test this assumption in two members of the Extant Phylogenetic Bracket of the Dinosauria, the emu, Dromaius novaehollandiae and the American alligator, Alligator mississippiensis, by investigating the sequence and timing of sutural fusion in their skulls. As expected, almost all the sutures in the emu skull progressively close (i.e., they get narrower) and then obliterate during ontogeny. However, in the American alligator, only two sutures out of 36 obliterate completely and they do so during embryonic development. Surprisingly, as maturity progresses, many sutures of alligators become wider in large individuals compared to younger, smaller individuals. Histological and histomorphometric analyses on two sutures and one synchondrosis in an ontogenetic series of American alligator confirmed our morphological observations. This pattern of sutural widening might reflect feeding biomechanics and dietary changes through ontogeny. Our findings show that progressive sutural closure is not always observed in extant archosaurs, and therefore suggest that cranial sutural fusion is an ambiguous proxy for assessing maturity in non-avian dinosaurs.     

小叶锦鸡儿天然居群叶形态性状变异研究

为揭示小叶锦鸡儿(Caragana microphylla)天然居群叶形态性状的变异规律及其生态适应性特征,该研究以10个小叶锦鸡儿天然居群为对象,通过多重比较、巢式方差分析、相关性分析、聚类分析和主成分分析等方法,对7个叶形态性状进行分析。结果表明：(1)小叶锦鸡儿叶形态性状在居群内和居群间均存在极显著差异(P < 0.01),平均变异系数为10.13%,不同性状的变异幅度为6.23%~12.78%;平均叶形态性状的表型分化系数为43.62%,居群内变异(30.09%)大于居群间变异(24.91%),说明居群内是其叶形态性状变异的主要来源。(2)相关性分析表明,环境因子对小叶锦鸡儿的叶形态性状变异有很大的影响,在地理空间上主要呈现出沿海拔梯度的变异模式;主成分分析的结果显示,小叶宽、叶柄宽和叶柄长对小叶锦鸡儿叶形态变异起主导作用;利用欧式距离对小叶锦鸡儿居群进行UPGMA聚类分析结果显示,基于叶形态性状和环境因子可分别将小叶锦鸡儿10个居群分为3类和2类,Mantel检验结果表明,小叶锦鸡儿的叶形态性状变异不存在地理连续性。研究结果为小叶锦鸡儿的适应性进化和开发利用提供了理论依据。     

A late-surviving basal theropod dinosaur from the latest Triassic of North America

The oldest theropod dinosaurs are known from the Carnian of Argentina and Brazil. However, the evolutionary diversification of this group after its initial radiation but prior to the Triassic-Jurassic boundary is still poorly understood because of a sparse fossil record near that boundary. Here, we report on a new basal theropod, Daemonosaurus chauliodus gen. et sp. nov., from the latest Triassic 'siltstone member' of the Chinle Formation of the Coelophysis Quarry at Ghost Ranch, New Mexico. Based on a comprehensive phylogenetic analysis, Daemonosaurus is more closely related to coeval neotheropods (e.g. Coelophysis bauri) than to Herrerasauridae and Eoraptor. The skeletal structure of Daemonosaurus and the recently discovered Tawa bridge a morphological gap between Eoraptor and Herrerasauridae on one hand and neotheropods on the other, providing additional support for the theropod affinities of both Eoraptor and Herrerasauridae and demonstrating that lineages from the initial radiation of Dinosauria persisted until the end of the Triassic. Various features of the skull of Daemonosaurus, including the procumbent dentary and premaxillary teeth and greatly enlarged premaxillary and anterior maxillary teeth, clearly set this taxon apart from coeval neotheropods and demonstrate unexpected disparity in cranial shape among theropod dinosaurs just prior to the end of the Triassic.     

Phylogeny of Recent Canidae (Mammalia, Carnivora): relative reliability and utility of morphological and molecular datasets

Zrzavý, J. & ?i?ánková, V. (2004). Phylogeny of Recent Canidae (Mammalia, Carnivora): relative reliability and utility of morphological and molecular datasets. — Zoologica Scripta, 33, 311–333. Phylogenetic relationships within the Canidae are examined, based on three genes (cytb, COI, COII) and 188 morphological, developmental, behavioural and cytogenetic characters. Both separate and combined phylogenetic analyses were performed. To inspect the phylogenetic ‘behaviour’ of individual taxa, basic phylogenetic analysis was followed by experimental cladistic analyses based on different data‐partition combinations and taxon‐removal analyses. The following phylogeny of the Recent Canidae is preferred: (1) Urocyon is the most basal canid; (2) Vulpes is a monophyletic genus (including Fennecus and Alopex); (3) the doglike canids (DC) form a clade (=Dusicyon + Pseudalopex + Lycalopex + Cerdocyon + Atelocynus + Chrysocyon + Speothos + Lycaon + Cuon + Canis), split into two subclades, South American and Afro‐Holarctic, with uncertain position of the Chrysocyon + Speothos subclade; (4) Canis is paraphyletic due to the position of Lycaon and Cuon. Otocyon and Nyctereutes are the most problematic canid genera, causing an unresolved branching pattern of Otocyon, Vulpes, Nyctereutes and DC clades. Reclassification of the two basal species of ‘Canis’ into separate genera is proposed (Schaeffia for ‘C.’ adustus, Lupulella for ‘C.’ mesomelas). Although the morphological dataset ranked poorly in both separate and simultaneous analyses (measured by number of minimum‐length topologies, relative number of resolved nodes in the strict consensus of all minimum‐length topologies, consistency and retention indices, nodal dataset influence, and number of extra steps required by the data partition to reach the topology of the combined tree), the morphological synapomorphies represent nearly one quarter of all synapomorphies in the combined tree. Among the hidden morphological support of the combined tree the developmental and behavioural characters are conspicuously abundant.     

Ontogenetic influence on neural spine bifurcation in diplodocoidea (dinosauria: Sauropoda): A critical phylogenetic character

Within Diplodocoidea (Dinosauria: Sauropoda), phylogenetic position of the three subclades Rebbachisauridae, Dicraeosauridae, and Diplodocidae is strongly influenced by a relatively small number of characters. Neural spine bifurcation, especially within the cervical vertebrae, is considered to be a derived character, with taxa that lack this feature regarded as relatively basal. Our analysis of dorsal and cervical vertebrae from small‐sized diplodocoids (representing at least 18 individuals) reveals that neural spine bifurcation is less well developed or absent in smaller specimens. New preparation of the roughly 200‐cm long diplodocid juvenile Sauriermuseum Aathal 0009 reveals simple nonbifurcated cervical neural spines, strongly reminiscent of more basal sauropods such as Omeisaurus. An identical pattern of ontogenetically linked bifurcation has also been observed in several specimens of the basal macronarian Camarasaurus, suggesting that this is characteristic of several clades of Sauropoda. We suggest that neural spine bifurcation performs a biomechanical function related to horizontal positioning of the neck that may become significant only at the onset of a larger body size, hence, its apparent absence or weaker development in smaller specimens. These results have significant implications for the taxonomy and phylogenetic position of taxa described from specimens of small body size. On the basis of shallow bifurcation of its cervical and dorsal neural spines, the small diplodocid Suuwassea is more parsimoniously interpreted as an immature specimen of an already recognized diplodocid taxon. Our findings emphasize the view that nonmature dinosaurs often exhibit morphologies more similar to their ancestral state and may therefore occupy a more basal position in phylogenetic analyses than would mature specimens of the same species. In light of this, we stress the need for phylogenetic reanalysis of sauropod clades where vital characters may be ontogenetically variable, particularly when data is derived from small individuals. J. Morphol., 2012. © 2012 Wiley Periodicals, Inc.     

Phylogenetic relationships within a patagonian clade of reptiles (Liolaemidae: Phymaturus) based on DNA sequences and morphology

Phymaturus is a clade of lizards that occurs at moderate to high elevations in western Argentina and the adjacent central region of Chile, as well as in various volcanic plateaus of the Patagonian region of Argentina. This genus had previously been divided into two groups: the patagonicus and the palluma groups. In this study, we analyzed relationships within the patagonicus group. The data set was built for 23 species plus nine other terminal taxa of undetermined taxonomic status. In total, 10,631 bp (ND4, Cytb, 12S, COI, five protein coding nuclear genes and seven anonymous nuclear loci) and 254 morphological characters were analyzed in a combined data set for 35 ingroup taxa and nine outgroups. We also ran separate DNA sequence and morphological data sets. We identified four main clades, and revealed congruencies and incongruences with previous studies. The indistinctus clade is recovered as the most basal within the patagonicus group in the strict parsimony analysis, while the somuncurensis clade is the most basal under Bayesian inference. The previously recovered calcogaster clade resulted paraphyletic in both analyses and part of their species are included in a redefined somuncurensis clade. We found low support at basal nodes provoked in part by contradictory evidence shown by rogue taxa. We show the phylogenetic information given by each partition/marker and how they contribute to relationships found in the total evidence analysis. We discuss the phylogenetic position of Phymaturus manuelae, Phymaturus tenebrosus, and Phymaturus patagonicus.     

The evolution of dinosaur tooth enamel microstructure

The evolution of tooth enamel microstructure in both extinct and extant mammalian groups has been extensively documented, but is poorly known in reptiles, including dinosaurs. Previous intensive sampling of dinosaur tooth enamel microstructure revealed that: (1) the three‐dimensional arrangement of enamel types and features within a tooth—the schmelzmuster—is most useful in diagnosing dinosaur clades at or around the family level; (2) enamel microstructure complexity is correlated with tooth morphology complexity and not necessarily with phylogenetic position; and (3) there is a large amount of homoplasy within Theropoda but much less within Ornithischia. In this study, the examination of the enamel microstructure of 28 additional dinosaur taxa fills in taxonomic gaps of previous studies and reinforces the aforementioned conclusions. Additionally, these new specimens reveal that within clades such as Sauropodomorpha, Neotheropoda, and Euornithopoda, the more basal taxa have simpler enamel that is a precursor to the more complex enamel of more derived taxa and that schmelzmusters evolve in a stepwise fashion. In the particularly well‐sampled clade of Euornithopoda, correlations between the evolution of dental and enamel characters could be drawn. The ancestral schmelzmuster for Genasauria remains ambiguous due to the dearth of basal ornithischian teeth available for study. These new specimens provide new insights into the evolution of tooth enamel microstructure in dinosaurs, emphasizing the importance of thorough sampling within broadly inclusive clades, especially among their more basal members.     

Evolution of heteroneuran Lepidoptera (Insecta) and the utility of dopa decarboxylase for Cretaceous-age phylogenetics

This study tests the utility of the nuclear gene encoding dopa decarboxylase (DDC) for recovering Cretaceous‐age divergences within the lepidopteran clade Heteroneura, which contains 98% of lepidopteran species. A 709‐bp fragment of DDC has been sequenced in 32 species, including representatives of all major lineages of Heteroneura plus outgroups from more basal lepidopteran groups and the related order Trichoptera. Pairwise divergences across the first and second codon positions and amino acids increase with depth throughout the taxonomic hierarchy, indicating that non‐synonymous substitutions are not fully saturated; whereas, divergences across the third codon position level off at the family to superfamily level. Inclusion of non‐neolepidopteran outgroups results in phylogeny estimates that contradict well established groups, almost surely due to sparse taxon sampling and high character divergence. When these taxa and an equivalently divergent basal ditrysian are excluded, DDC trees show nearly complete recovery of ten uncontroversial basal heteroneuran ‘test clades’ of family rank and higher, about half with strong bootstrap support. Thus, DDC clearly carries phylogenetic signal at these levels. Bootstrap support for resolution of the controversial relationships among the five main heteroneuran groups (four monotrysian superfamilies plus Ditrysia) is individually low, but two of three previous hypotheses were statistically rejected overall by DDC. DDC trees within the primitive heteroneuran superfamily Incurvarioidea, though modestly supported, closely resemble a previous morphological hypothesis, while removing the requirement for reversal in a possible ‘key adaptation’, the larval case. Taxon overlap with a previous mtDNA study of Prodoxidae (Incurvarioidea), which includes much‐ studied mutualist pollinators, permits a comparison of substitution rates with the conservative mitochondrial COI+COII region, as well as combined‐data re‐examination of generic reltionships. Non‐synonymous substitution is about 25% slower in DDC than in COI+COII, though synonymous substitution is faster. With additional taxon sampling, and in combination with other genes, DDC promises to be a powerful tool for reconstructing among‐superfamily relationships within Lepidoptera and probably other insect groups.     

The origins of Dinosauria: much ado about nothing

Research this century has greatly improved our knowledge of the origin and early radiation of dinosaurs. The unearthing of several new dinosaurs and close outgroups from Triassic rocks from various parts of the world, coupled with improved phylogenetic analyses, has set a basic framework in terms of timing of events and macroevolutionary patterns. However, important parts of the early dinosauromorph evolutionary history are still poorly understood, rendering uncertain the phylogenetic position of silesaurids as either non‐dinosaur Dinosauriformes or ornithischians, as well as that of various early saurischians, such as Eoraptor lunensis and herrerasaurs, as either noneusaurischians or members of the sauropodomorph or theropod lineages. This lack of agreement in part derives from a patchy distribution of traits among early members of the main dinosauromorph lineages and requires a more meticulous assessment of characters and homologies than those recently conducted. Presently, the oldest uncontroversial dinosaur records come from Late Triassic (Carnian) rocks of South America, southern Africa and India, hinting at a south‐western Pangaea origin of the group. Besides, macroevolutionary approaches suggest that the rise of dinosaurs was a more gradual process than previously understood. Obviously, these tentative scenarios need to be tested by new fossil finds, which should also help close the major gaps recognized in the fossil record of Triassic dinosauromorphs.     

The phylogeny of Sericini and their position within the Scarabaeidae based on morphological characters (Coleoptera: Scarabaeidae)

Abstract. To reconstruct the phylogeny of the Sericini and their systematic position among the scarabaeid beetles, cladistic analyses were performed using 107 morphological characters from the adults and larvae of forty‐nine extant scarabaeid genera. Taxa represent most ‘traditional’ subfamilies of coprophagous and phytophagous Scarabaeidae, with emphasis on the Sericini and other melolonthine lineages. Several poorly studied exoskeletal features have been examined, including the elytral base, posterior wing venation, mouth parts, endosternites, coxal articulation, and genitalia. The results of the analysis strongly support the monophyly of the ‘orphnine group’ + ‘melolonthine group’ including phytophagous scarabs such as Dynastinae, Hopliinae, Melolonthinae, Rutelinae, and Cetoniinae. This clade was identified as the sister group to the ‘dung beetle line’ represented by Aphodius + Copris. The ‘melolonthine group’ is comprised in the strict consensus tree by two major clades and two minor lineages, with the included taxa of Euchirinae, Rutelinae, and Dynastinae nested together in one of the major clades (‘melolonthine group I’). Melolonthini, Cetoniinae, and Rutelinae are strongly supported, whereas Melolonthinae and Pachydemini appear to be paraphyletic. Sericini + Ablaberini were identified to be sister taxa nested within the second major melolonthine clade (‘melolonthine group II’). As this clade is distributed primarily in the southern continents, one could assume that Sericini + Ablaberini are derived from a southern lineage. Plausibly, ancestors of Sericini + Ablaberini and Athlia were separated by a vicariance event, such as the separation of the African plate from the rest of Gondwana, whereas Sericini and Ablaberini probably diversified during the early Tertiary, with dispersal of some basal Sericini to South America.     

A New Mass Mortality of Juvenile Protoceratops and Size-Segregated Aggregation Behaviour in Juvenile Non-Avian Dinosaurs

Background

Monodominant bonebeds are a relatively common occurrence for non-avian dinosaurs, and have been used to infer associative, and potentially genuinely social, behavior. Previously known assemblages are characterized as either mixed size-classes (juvenile and adult-sized specimens together) or single size-classes of individuals (only juveniles or only adult-sized individuals within the assemblage). In the latter case, it is generally unknown if these kinds of size-segregated aggregations characterize only a particular size stage or represent aggregations that happened at all size stages. Ceratopsians (“horned dinosaurs”) are known from both types of assemblages.

Methods/Principal Findings

Here we describe a new specimen of the ceratopsian dinosaur Protoceratops andrewsi, Granger and Gregory 1923 from Mongolia representing an aggregation of four mid-sized juvenile animals. In conjunction with existing specimens of groups of P. andrewsi that includes size-clustered aggregations of young juveniles and adult-sized specimens, this new material provides evidence for some degree of size-clustered aggregation behaviour in Protoceratops throughout ontogeny. This continuity of size-segregated (and presumably age-clustered) aggregation is previously undocumented in non-avian dinosaurs.

Conclusions

The juvenile group fills a key gap in the available information on aggregations in younger ceratopsians. Although we support the general hypothesis that many non-avian dinosaurs were gregarious and even social animals, we caution that evidence for sociality has been overstated and advocate a more conservative interpretation of some data of ‘sociality’ in dinosaurs.     

Galvechelone lopezmartinezae gen. et sp. nov., a new cryptodiran turtle in the Lower Cretaceous of Europe

Abstract: The Spanish town of Galve (Teruel) is notable because of the abundance of Upper Jurassic and, especially, Lower Cretaceous vertebrates recorded there. Although most groups have been studied in detail, information on turtles is very limited even though the material is relatively abundant. So far, no turtle taxa have been identified at the generic level. The only Lower Cretaceous articulated specimen from Galve is analysed here. It is identified as a representative of Cryptodira, Galvechelone lopezmartinezae gen. et sp. nov. Galvechelone lopezmartinezae is determined as a taxon belonging to the node that groups the turtles traditionally assigned to ‘Macrobaenidae’ and ‘Sinemydidae’, and other taxa such as the members of Panchelonioidea. This node, very abundant in the Lower Cretaceous of Asia, and with a broad subsequent distribution, has recently been recognized in the Lower Cretaceous of Europe. The diversity of basal members of Eucryptodira in the European Late Jurassic (represented by Thalassemydidae, Plesiochelyidae and Eurysternidae) was high. Owing to a relative scarcity of well‐preserved early Cretaceous turtles from Europe, the knowledge of this group of reptiles is limited. The study of the new turtle from Galve, together with the recently described Hoyasemys jimenezi, and the recently completed review of the enigmatic Chitracephalus dumonii demonstrate that members of the cryptodiran node grouping ‘Macrobaenidae’, ‘Sinemydidae’ and Panchelonioidea were also very diverse in this period.     

A Family Level Analysis of Tardigrade Phylogeny

In the present study a character data set suitable for cladistic analysis at the family level was developed. A data matrix consisting of 50 morphological characters from 15 families of tardigrades was analyzed by maximum parsimony. Kinorhynchs, loriciferans, and gastrotrichs were used as outgroups. The results agree with the currently accepted hypothesis that Eutardigrada and Heterotardigrada are distinct monophyletic groups. Among the eutardigrades, Eohypsibiidae was found to be a sister group to Macrobiotidae+Hypsibiidae, while Milnesiidae was the basal eutardigrade family. The basal heterotardigrade family was found to be Oreellidae. Echiniscoideans grouped with some traditional Arthrotardigrada (Renaudarctidae, Coronarctidae+Batillipedidae) suggesting that the arthrotardigrades are not monophyletic. The 18S rRNA gene sequence of Batillipes mirus Richters, 1909 and Calohypsibius schusteri Nelson & McGlothlin, 1996 were obtained and their addition to a previously published dataset supports the monophyly of Heterotardigrada and of Parachela versus Apochela within the Eutardigrada.