Skull ontogeny in Arrhinoceratops brachyops (Ornithischia: Ceratopsidae) and other horned dinosaurs

Disentangling ontogenetic from interspecific variation is key to understanding biodiversity in the fossil record, yet information on growth in the ceratopsid subfamily Chasmosaurinae is sparse. Here, we describe the partial skull of a juvenile chasmosaurine, attributed to Arrhinoceratops brachyops, within the context of more mature specimens of this species, to better understand the ontogenetic transformations therein. We show that as A. brachyops matured, the postorbital horncores became longer and shifted from a posterior to an anterior inclination, the delta‐shaped frill epiossifications became lower and fused to the underlying frill, and the face became more elongate. In these respects, A. brachyops closely resembled Triceratops, suggesting that these ontogenetic changes may have been common to all long‐horned chasmosaurines. However, an event‐paired cladistic analysis of Chasmosaurinae using a standardized matrix of 24 developmental characters reveals that the relative timing of ontogenetic events in Arrhinoceratops was more like that of Chasmosaurus, particularly in the relatively late reduction in scalloping around the frill margins. Thus, the ontogenetic similarities between Arrhinoceratops and Triceratops appear to be plesiomorphic, partly related to the retention of the elongate postorbital horncores, which are primitive for Ceratopsidae. This study elucidates the otherwise contentious evolutionary relationships of Arrhinoceratops, and highlights the importance of ontogenetic data for resolving phylogenies when morphological data from adults alone are inadequate. © 2015 The Linnean Society of London     

Is Torosaurus Triceratops? Geometric Morphometric Evidence of Late Maastrichtian Ceratopsid Dinosaurs

Background

Recent assessments of morphological changes in the frill during ontogeny hypothesized that the late Maastrichtian horned dinosaur Torosaurus represents the “old adult” of Triceratops, although acceptance of this finding has been disputed on several lines of evidence.

Methodology/Principal Findings

Examining the cranial morphology of 28 skulls in lateral view and 36 squamosals of Nedoceratops hatcheri, Triceratops spp. and Torosaurus spp. by means of landmark-based geometric morphometrics, we compared ontogenetic trajectories among these taxa. Principal Component Analysis and cluster analysis confirmed different cranial morphologies. Torosaurus shape space is well separated from Triceratops, whereas Triceratops horridus and Triceratops prorsus partially overlap within Triceratops shape space. Linear regressions between shape and size suggest different ontogenetic trajectories among these taxa. Results support the “traditional” taxonomic status of Torosaurus. We hypothesize that ontogeny drives cranial morphology with different patterns between Torosaurus and Triceratops.

Conclusions/Significance

Torosaurus is a distinct and valid taxon. Whether looking at entire skulls, skulls without the frill, frills alone, or squamosals, Torosaurus has different morphologies and distinct allometric trajectories compared to Triceratops. This new approach confirms the taxonomic status of Torosaurus as well as the comparatively low diversity of ceratopsids at the end of the Maastrichtian in North America.     

Dental structure of the Giant lantern shark <Emphasis Type="Italic">Etmopterus baxteri</Emphasis> (Chondrichthyes: Squaliformes) and its taxonomic implications

Dogfish sharks (Squaliformes) are a highly diverse group of neoselachians occurring in a wide range of marine environments and are common members of deep-sea faunas. The order Squaliformes comprises six families with approximately 98 extant species. The dentition of most squaliforms is characterized by a strong dignathic heterodonty and dental variation yielding a suite of potential tooth characters that could be used for taxonomic and systematic purposes. So far, no detailed study has been carried out to analyse the use of tooth morphologies in reconstructing the phylogeny of squaliforms. Also, the degree of characteristics of intraspecific variability of tooth morphologies is still unclear. Here, we analysed the dental differences between juveniles and adults and between the sexes of the Giant lantern shark, Etmopterus baxteri, and tested these dental characters for taxonomic purposes employing different statistical procedures. The results show that upper teeth of adult females and males differ morphologically in that those of females are bigger and display a lanceolate central cusp, whereas male specimens have thin and needle-like central cusps. Upper teeth of males have a higher number and a more pronounced variability of lateral cusplets than those of females. Moreover, an ontogenetic heterodonty might be developed in male specimens with sexually immature males displaying similar dental morphologies to those of adult females. Lower teeth, conversely, do not differ morphologically between the sexes. Results indicate that tooth morphologies of squaliform sharks bear high potential for phylogenetic purposes if tooth variations are considered, but have to be treated with care, if no variation is analysed.     

Morphological and molecular investigation of polymorphism and cryptic species in tanaid crustaceans: implications for tanaid systematics and biodiversity estimates

A combination of traditional taxonomic procedures and molecular techniques has provided new insight into the problems of cryptic species and sexual and ontogenetic polymorphism in the Tanaidacea. Using polymerase chain reaction and DNA markers, three cryptic species of Paratanais were identified. PCR primers were used to amplify the divergent internal transcribed spacers (ITS) of these species. Restriction digestion of the amplified rDNA generated species specific DNA banding. Male and five female stages of Paratanais maleticus sp. nov. and two other new species, P. malign us and P. perturbatius , are described. Morphological variation, both sexual and ontogenetic, was found in several generic characters of Paratanais and required the diagnosis to be modified. The identification of three undescribed cryptic species from a single microhabitat in a well studied, shallow water and easily accessible locality, demonstrate that the biodiversity of tanaid crustacean is significantly underestimated.     

Quantitative handling of characters useful in snake systematics with particular reference to intraspecific variation in the Ringed Snake Natrix natrix (L.)

This paper is based on the numerous characters used in a biometric study of the intraspecific variation of Natrix natrix (L.). These characters are drawn from the internal morphology, scalation, colour pattern, body proportions, dentition and dermal sense organs. The criteria involved in selecting these characters for taxonomic work are considered.
The alternative methods for quantifying the above features are discussed and so are the theoretical problems involved in coding the continuous characters.
The value of the characters in studying the geographic variation, ontogenetic variation and sexual dimorphism in N. natrix is indicated.
The use of a wider range of characters than just the scalation and colour pattern is urged when diagnosing the lower taxonomic levels of ophidians. The neglected internal morphology may well be able to supply characters to this end.     

Ontogenetic shifts in functional morphology of dragonfly legs (Odonata: Anisoptera)

Anisopteran leg functions change dramatically from the final larval stadium to the adult. Larvae use legs mainly for locomotion, walking, climbing, clinging, or burrowing. Adults use them for foraging and grasping mates, for perching, clinging to the vegetation, and for repelling rivals. In order to estimate the ontogenetic shift in the leg construction from the larva to the adult, this study quantitatively compared lengths of fore, mid, and hind legs and the relationships between three leg segments, femur, tibia, and tarsus, in larval and adult Anisoptera of the families Gomphidae, Aeshnidae, Cordulegastridae, Corduliidae, and Libellulidae, represented by two species each. We found that leg segment length ratio as well as ontogenetic shift in length ratios was different between families, but rather similar within the families. While little ontogenetic shift occurred in Aeshnidae, there were some modifications in Corduliidae and Libellulidae. The severest shift occurred in Gomphidae and Cordulegastridae, both having burrowing larvae. These two families form a cluster, which is in contrast to their taxonomic relationship within the Anisoptera. Cluster analysis implies that the function of larval legs is primarily responsible for grouping, whereas adult behavior or the taxonomic relationships do not explain the grouping. This result supports the previous hypothesis about the convergent functional shift of leg characters in the dragonfly ontogenesis.     

The use of pigmentation and morphological characters to identify Italian mullet fry

Mullet fry from Italian coastal waters are examined to ascertain ontogenetic changes in pigmentation, external morphology, anal-fin ray counts, and morphology of the pyloric caeca and pharyngobranchial organ. The taxonomic utility of these characters is assessed. Intraspecific and ontogenetic variation in anal-fin ray numbers do not make this a consistently reliable character for identification purposes. Pharyngobranchial morphology can assist diagnoses of larger fry or juveniles; however, pigmentation patterns are the most reliable characters for smaller fry. Pigmentation characters, which can be used in laboratory- and field-based studies, require minimal manipulation of the fish and enable rapid identification of large numbers of specimens. These techniques are valuable for aquaculturists and fish biologists requiring more information on the identification, distribution and life history of mugilids.     

A remarkable short-snouted horned dinosaur from the Late Cretaceous (late Campanian) of southern Laramidia

The fossil record of centrosaurine ceratopsids is largely restricted to the northern region of western North America (Alberta, Montana and Alaska). Exceptions consist of single taxa from Utah (Diabloceratops) and China (Sinoceratops), plus otherwise fragmentary remains from the southern Western Interior of North America. Here, we describe a remarkable new taxon, Nasutoceratops titusi n. gen. et sp., from the late Campanian Kaiparowits Formation of Utah, represented by multiple specimens, including a nearly complete skull and partial postcranial skeleton. Autapomorphies include an enlarged narial region, pneumatic nasal ornamentation, abbreviated snout and elongate, rostrolaterally directed supraorbital horncores. The subrectangular parietosquamosal frill is relatively unadorned and broadest in the mid-region. A phylogenetic analysis indicates that Nasutoceratops is the sister taxon to Avaceratops, and that a previously unknown subclade of centrosaurines branched off early in the group''s history and persisted for several million years during the late Campanian. As the first well-represented southern centrosaurine comparable in age to the bulk of northern forms, Nasutoceratops provides strong support for the provincialism hypothesis, which posits that Laramidia—the western landmass formed by inundation of the central region of North America by the Western Interior Seaway—hosted at least two coeval dinosaur communities for over a million years of late Campanian time.     

Postnatal Cranial Development in Papionin Primates: An Alternative Model for Hominin Evolutionary Development

The evolution of hominin growth and life history has long been a subject of intensive research, but it is only recently that paleoanthropologists have considered the ontogenetic basis of human morphological evolution. To date, most human EvoDevo studies have focused on developmental patterns in extant African apes and humans. However, the Old World monkey tribe Papionini, a diverse clade whose members resemble hominins in their ecology and population structure, has been proposed as an alternative model for human craniofacial evolution. This paper reviews prior studies of papionin development and socioecology and presents new analyses of juvenile shape variation and ontogeny to address fundamental questions concerning primate cranial development, including: (1) When are cranial shape differences between species established? (2) How do epigenetic influences modulate early-arising pattern differences? (3) How much do postnatal developmental trajectories vary? (4) What is the impact of developmental variation on adult cranial shape? and, (5) What role do environmental factors play in establishing adult cranial form? Results of this inquiry suggest that species differences in cranial morphology arise during prenatal or earliest postnatal development. This is true even for late-arising features that develop under the influence of epigenetic factors such as mechanical loading. Papionins largely retain a shared, ancestral pattern of ontogenetic shape change, but large size and sexual dimorphism are associated with divergent developmental trajectories, suggesting differences in cranial integration. Developmental simulation studies indicate that postnatal ontogenetic variation has a limited influence on adult cranial morphology, leaving early morphogenesis as the primary determinant of cranial shape. The ability of social factors to influence craniofacial development in Mandrillus suggests a possible role for phentotypic plasticity in the diversification of primate cranial form. The implications of these findings for taxonomic attribution of juvenile fossils, the developmental basis of early hominin characters, and hominin cranial diversity are discussed.     

Etude du répertoire vocal du Gibbon a favoris blancs (Hylobates concolor leucogenys): Analyse structurale des vocalisations

The basic vocal repertoire of the Crested Gibbon (Hylobates concolor leucogenys) can be defined from qualitative and quantitative analysis of the vocalizations. From sonagraphic analysis of calls of 9 juveniles, one adult ♂, one sub-adult ♂ and one sub-adult ♀ arranged in three semi-free living groups, one can describe six vocal types on a morphological basis. The sound repertoire of juveniles is composed of graded signals which seem to become more discrete and fewer as the individual matures. These features are compared with other Cercopithecides and Pongides on the basis of eco-ethological and ontogenetic implications.     

Geographical variation in size in the European rabbit Oryctolagus cuniculus (Lagomorpha: Leporidae) in western Europe and North Africa

Craniometric data of nine characters from 260 skulls of Oryctolagus cuniculus from western Europe and northern Africa were gathered to examine phenotypic variation in size, using both univariate and multivariate analyses. Although animals from the northern part of Europe were found to be larger than those from southern Europe, variation in size occurred in a continuous cline. Skull size was positively correlated with latitude and negatively correlated with ambient temperature. On the basis of skull morphology, there is no evidence for separation of O. cuniculus into the two or more subspecies commonly described. Further data, particularly from genetic material, are required to clarify the taxonomic status of the rabbit in Europe.     

Ontogenetic systematics: The synthesis of taxonomy, phylogenetics, and evolutionary developmental biology

The main purpose of the present review is to draw attention to growing problems in the modern systematics and phylogenetics which are presently underestimated by the professional community. The dramatic reduction of the importance of ontogeny and morphology in phylogenetic studies of the second part of the 20th century is considered among the major factors of the modern taxonomic and evolutionary paradigm. The deep contradiction of modern approaches, which either merely consider systematics and phylogeny as genealogy or even in a neotypolgical manner irrespective of the evolutionary idea, is demonstrated. Thus, despite the widespread opinion that the evolutionary theory is the major basis for taxonomy, the processes, which in fact caused the origin and formation of the systematic hierarchy are often considered as redundant for the procedure of classification. In this respect, the classical, but well forgotten statement that evolution is a modification of ontogeny is specially highlighted. Tight relationships between evolution, ontogeny, systematics, and phylogenetics are prima facie obvious, but also presently underestimated, although the field of the evo-devo is continuously growing. Paradoxically, even despite the outburst of various molecular ontogenetic approaches, the commonly accepted evolutionary paradigm still lacks a general theory for changes in the shape of organisms. As a step towards the development of such a theory, a synthesis (or more exactly, resynthesis) of still largely independently developing major biological fields, i.e., ontogenetic and evolutionary studies, on the one hand, and traditional taxonomy, on the other hand, a new concept of ontogenetic systematics is proposed. The new concept is intended for integration of supposedly ??immobile?? traditional taxonomy with the dynamics, but predominantly considered as hypothetical, evolutionary field based on the process of ontogeny, which, in contrast to the evolution itself, can be observed in the real time. Therefore, it is concluded that, for instance, the evolution of the main group of living organisms Metazoa, is primarily the evolution of a very limited number of ontogenetic cycles that were formed as early as the Early Cambrian. A significant underestimation of cyclic properties of ontogeny in the evolution and systematics is shown. Using two model groups, echinoderms of the class Ophiuroidea and dorid nudibranch mollusks (Gastropoda: Doridacea), practical importance of the integrative approach developed here is demonstrated. The ??disruption?? of the ancestral ontogenetic cycle and further formation of a new descendant cycle (which implies some continuity of ancestral and descendant characters) is considered to be a major evolutionary pattern. The model proposed implies either progressive (addition of stages and characters) or regressive (reduction of already existing stages and structures) modification of ancestral taxon, the diagnosis of which corresponds to the model of its ontogenetic cycle. In the extreme cases of disruption of the ancestral ontogenetic cycle, adult characters of descendants are substituted by juvenile ancestral features, demonstrating paedomorphoses in the narrow sense. Within the framework of the approach proposed, the evolutionary and ontogenetic models of ancestral ontogenetic cycles of brittle stars and dorid nudibranchs are developed and discussed. Based on the original material of the extinct Paleozoic ophiuroid group Oegophiurida, the origin of key evolutionary novelties is discussed. A major conclusion of the present review is the high necessity of integration of new molecular data with already well-established taxonomic hierarchy and ontogenetic information as a basis for the development of the general theory of transformations of organisms, i.e., the theory of evolution in its true sense.     

雄性大熊猫对化学信息行为反应的年龄差异

化学通讯对哺乳动物的生存和繁殖起着重要作用。研究了雄性大熊猫(Ailuropoda melanoleuca)对同伴个体尿液气味行为反应的发育模式。结果显示,在成年雌性个体的尿液气味刺激下,雄性个体表现显著多的嗅闻行为和嗅闻/舔舐环境行为,但是嘶咬气味刺激物的行为明显减少。在雌性个体的尿液气味刺激下,不同年龄段的雄性个体行为表现不同,成年雄性个体表现较亚成年和幼年个体显著多的舔舐行为。此外,成年个体和亚成年个体均表现较多的嗅闻/舔舐环境行为,而幼年个体则无该行为表现。幼年个体较成年和亚成年个体表现显著多的气味涂抹行为,而且嘶咬气味刺激物的时间较亚成年个体显著多。幼年个体和亚成年个体对雌性和雄性个体尿液气味刺激的行为反应不存在显著差异。研究结果表明,雄性大熊猫对同种个体尿液中化学信息的行为反应呈现出年龄差异。     

Early development of the facial region in a non-avian theropod dinosaur

An isolated maxilla of the theropod dinosaur Allosaurus from the Late Jurassic (the Kimmeridgian, 153 million years ago) of Portugal is the first cranial remain of a non-coelurosaurian theropod hatchling reported so far, and sheds new light on the early cranial development of non-avian theropods. Allosaurus hatchlings seem to have been one-seventh or less of the adult length and are thus comparable in relative size to hatchlings of large extant crocodile species, but are unlike the relatively larger hatchlings in coelurosaurs. The snout experienced considerable positive allometry and an increase in tooth count during early development. The element is especially noteworthy for the abundant and well-developed features associated with the paranasal pneumatic system. Pneumatic structures present include all those found in adult allosaurids and most are even more developed than in adult skulls. Together with evidence on the ontogeny of the tympanic pneumatic system in allosaurids, these findings demonstrate that cranial pneumaticity developed early in theropod ontogeny. The strong development of pneumatic features in early ontogenetic stages of non-avian theropods supports the hypothesis that pneumatization of cranial bones was opportunistic and indicates that heterochrony played an important role in the evolution of craniofacial pneumaticity in this group.     

Evolution of ontogeny in the hippopotamus skull: using allometry to dissect developmental change

Allometry describes the effect of size change on aspects of an organism's form and can be used to summarize the developmental history of growing parts of an animal. By comparing how allometric growth differs between species, it is possible to reveal differences in their pathways of development. The ability to compare and categorize developmental change between species is demonstrated here using morphometric methods. This involves the interspecific statistical comparison of a large number of bivariate relationships that summarize ontogenetic trajectories. These linear ontogenetic trajectories can be modified as they evolve in any of three ways: ontogenetic scaling indicative of change in the duration of growth, lateral shifts indicative of changes in prenatal development, and directional change indicative of novel modes of postnatal growth. I apply this analysis to skulls of the common hippopotamus ( Hippopotamus amphibius ) and the pygmy hippopotamus ( Hexaprotodon liberiensis ). The number of allometric changes falling into each category was statistically determined and Jolicoeur's multivariate generalization of simple allometry was used to provide an overview of cranial variation. For these skulls, directional change was not found to be statistically significant, but ontogenetic scaling and lateral shifts were both common. This indicates that conserved patterns of growth covariance (ontogenetic scaling) can be separated from novel or derived patterns (directional change and/or lateral shifts). This study demonstrates that He. liberiensis is not simply an ontogenetically scaled version of its larger relative. The evolutionary implications of allometric growth variation are discussed in the light of these findings and those of other studies.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 625–638.     

An evaluation of the systematic value of skull morphology in the Trimeresurus radiation (Serpentes: Viperidae: Crotalinae) of Asian pitvipers

Trimeresurus (in its widest sense) is a very diverse and widespread radiation of Asian pitvipers, which has been subject to numerous taxonomic revisions, some of which have been based on characteristics of the skull. In order to evaluate the taxonomic utility of such characters, we conducted a comparison of the skulls of 57 specimens representing nine genera, and two currently unassigned species that are very closely related to each other. A canonical variate analysis reveals three distinct phenetic groups: the Protobothrops group, a group containing Ovophis monticola , ' Ovophis ' okinavensis , and ' Trimeresurus ' gracilis , and finally a group comprising the remaining species and characterized by considerable overlap between most genera with the exception of the monotypic Peltopelor and Himalayophis . Agreement between phenetic similarity based on skull characteristics and phylogenetic relationships based on molecular evidence varies between different groups: the morphological similarity of the skull of Protobothrops sieversorum to the other Protobothrops species is congruent with their recent synonymization while the phenetic similarity among the species within the second group does not reflect current molecular phylogenetic relationships and indicates that convergent or parallel evolution may be responsible for at least some of the phenetic similarity detected among skulls of the Asian pit vipers examined. A test of phylogenetic independence, however, indicates that there is still a significant phylogenetic signal that can be recovered from several skull characteristics. Thus, we conclude that skull morphology can contribute to an overall understanding of pitviper taxonomy, but that it would be unwise to rely on skull characteristics alone.     

Ontogeny of robusticity of craniofacial traits in modern humans: A study of South American populations

To date, differences in craniofacial robusticity among modern and fossil humans have been primarily addressed by analyzing adult individuals; thus, the developmental basis of such differentiation remains poorly understood. This article aims to analyze the ontogenetic development of craniofacial robusticity in human populations from South America. Geometric morphometric methods were used to describe cranial traits in lateral view by using landmarks and semilandmarks. We compare the patterns of variation among populations obtained with subadults and adults to determine whether population‐specific differences are evident at early postnatal ontogeny, compare ontogenetic allometric trajectories to ascertain whether changes in the ontogeny of shape contribute to the differentiation of adult morphologies, and estimate the amount of size change that occurs during growth along each population‐specific trajectory. The results obtained indicate that the pattern of interpopulation variation in shape and size is already established at the age of 5 years, meaning that processes acting early during ontogeny contribute to the adult variation. The ontogenetic allometric trajectories are not parallel among all samples, suggesting the divergence in the size‐related shape changes. Finally, the extension of ontogenetic trajectories also seems to contribute to shape variation observed among adults. Am J Phys Anthropol 2010. © 2009 Wiley‐Liss, Inc.