Cranial morphology of the plagiosaurid Gerrothorax pulcherrimus as an extreme example of evolutionary stasis

Schoch, R.R. & Witzmann, F. 2011: Cranial morphology of the plagiosaurid Gerrothorax pulcherrimus as an extreme example of evolutionary stasis. Lethaia, Vol. 45, pp. 371–385. The plagiosaurid Gerrothorax pulcherrimus from the Triassic of Greenland and Germany is represented by skulls ranging from 4 to 12 cm in length and sheds light on ontogeny, individual variation, and variation in time and space. Ontogeny was remarkably stable in G. pulcherrimus, with the smallest known specimens resembling the adults closely in most features. A true ontogenetic change is evident in the ornament of dermal bones, in that the smallest specimens have ridges whereas in the successively larger ones, pustules spread over increasingly larger areas. The skull becomes proportionally longer, and the adductor chambers relatively narrower. The positive allometry of both the orbits and the interpterygoid vacuities suggests that the eye supporting musculature – rather than the jaw adductors – increased proportionally during growth. Individual, not age‐related variation in the dermal skull roof affects partial fusion of parietals, presence and extent of the interfrontoparietal, and the morphological pattern of the posterior skull table. The ventral surface of the basal plate of the parasphenoid ranges from smooth over poorly to heavily ornamented or dentigerous. Considering the impressive longevity of more than 35 Myr, the morphological changes of G. pulcherrimus are minor. Our ecological interpretation for G. pulcherrimus is that it relied on the permanent presence of water, but was flexible with respect to the size and nature of the water body as well as to changes in salinity. The unparalleled extent of evolutionary stasis may therefore be based on the ecological flexibility of this morphologically so tightly constrained temnospondyl. □Ecological flexibility, ontogeny,Temnospondyli, Triassic, variation.     

The plagiosaurid temnospondyl Plagiosuchus pustuliferus (Amphibia: Temnospondyli) from the Middle Triassic of Germany: anatomy and functional morphology of the skull

The cranial anatomy of the plagiosaurid temnospondyl Plagiosuchus pustuliferus, from the Middle Triassic of Germany, is described in detail on the basis of a newly discovered skull and mandibular material. The highly derived skull is characterized by huge orbitotemporal fenestrae, a reduction of the circumorbital bones – the prefrontal, postfrontal and (probably) postorbital are lost – and the expansion of the jugal to occupy most of the lateral skull margin. Ventrally the extremely long subtemporal vacuities correlate with the elongate adductor fossa of the mandible. The dentition is feebly developed on both skull and mandible. Ossified ?ceratobranchials and ‘branchial denticles’ indicate the presence of open gills clefts in life. The remarkably divergent cranial morphology of P. pustuliferus highlights the extraordinary cranial diversity within the Plagiosauridae, probably unsurpassed within the Temnospondyli. Specific structural aspects of the skull – including an extremely short marginal tooth row, feeble dentition and an elongated chamber for adductor musculature – together with evidence for a hyobranchial skeleton, suggests that P. pustuliferus utilized directed suction feeding for prey capture. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 348–373.     

Braincase,palatoquadrate and ear region of the plagiosaurid Gerrothorax pulcherrimus from the Middle Triassic of Germany

Abstract: The complete neurocranium plus palatoquadrate of the plagiosaurid temnospondyl Gerrothorax pulcherrimus from the Middle Triassic of Germany is described for the first time, based on outer morphological observations and micro‐CT scanning. The exoccipitals are strong elements with paroccipital processes and well‐separated occipital condyles. Anterolaterally, the exoccipitals contact the otics, which are mediolaterally elongated and have massive lateral walls. The otics contact the basisphenoid, which shows well‐developed sellar processes. Anteriorly, the basisphenoid is continuous with the sphenethmoid region. In its posterior portion, the sphenethmoid gives rise to robust, laterally directed laterosphenoid walls, a unique morphology among basal tetrapods. The palatoquadrate is extensively ossified. The quadrate portion overlaps the descending lamina of squamosal and ascending lamina of pterygoid anteriorly, almost contacting the epipterygoid laterally. The epipterygoid is a complex element and may be co‐ossified with otics and laterosphenoid walls. It has a broad, sheet‐like footplate and a horizontally aligned ascending process that contacts the laterosphenoid walls. The degree of ossification of the epipterygoid, however, is subject to individual variation obviously independent from ontogenetic changes. The stapes of Gerrothorax is a large, blade‐like element that differs conspicuously from the plesiomorphic temnospondyl condition. It has a prominent anterolateral projection which has not been observed in other basal tetrapods. Morphology of neurocranium and palatoquadratum of Gerrothorax most closely resembles that of the Russian plagiosaurid Plagiosternum danilovi, although the elements are less ossified in the latter. The extensive endocranial ossification of Gerrothorax is consistent with the general high degree of ossification in the exo‐ and endoskeleton of this temnospondyl and supports the view that a strong endocranial ossification cannot be evaluated as a plesiomorphic character in basal tetrapods.     

Mapping bone cell distributions to assess ontogenetic origin of primate midfacial form

Midfacial reduction in primates has been explained as a byproduct of other growth patterns, especially the convergent orbits. This is at once an evolutionary and developmental explanation for relatively short snouts in most modern primates. Here, we use histological sections of perinatal nonhuman primates (tamarin, tarsier, loris) to investigate how orbital morphology emerges during ontogeny in selected primates compared to another euarchontan (Tupaia glis). We annotated serial histological sections for location of osteoclasts or osteoblasts, and used these to create three‐dimensional “modeling maps” showing perinatal growth patterns of the facial skeleton. In addition, in one specimen we transferred annotations from histological sections to CT slices, to create a rotatable 3D volume that shows orbital modeling. Our findings suggest that growth in the competing orbital and neurocranial functional matrices differs among species, influencing modeling patterns. Distinctions among species are observed in the frontal bone, at a shared interface between the endocranial fossa and the orbit. The medial orbital wall is extensively resorptive in primates, whereas the medial orbit is generally depositional in Tupaia. As hypothesized, the orbital soft tissues encroach on available interorbital space. However, eye size cannot, by itself, explain the extent of reduction of the olfactory recess. In Loris, the posterior portion of medial orbit differed from the other primates. It showed evidence of outward drift where the olfactory bulb increased in cross‐sectional area. We suggest the olfactory bulbs are significant to orbit position in strepsirrhines, influencing an expanded interorbital breadth at early stages of development. Am J Phys Anthropol 154:424–435, 2014. © 2014 Wiley Periodicals, Inc.     

Effect of life history on the squirrel monkey (Platyrrhini,Saimiri) cranium

Among primates, squirrel monkeys uniquely possess an interorbital fenestra, in which the midline bony orbitosphenoid septum is largely absent and the soft tissues of the orbits are separated only by a thin membrane. Neural development may contribute to the approximation of the orbits to the midline in Saimiri, insofar as other platyrrhines with relatively large brains also have relatively narrow interorbital spaces compared to their near relatives. In Saimiri the narrow spacing of the orbits is further exacerbated by intense predation pressure on infants that may select for precocial neonates. The result is a large-headed neonate that is subject to unusual parturition constraints. These parturition constraints apply to the size and dolichocephalic shape of the squirrel monkey head in general, and to the relatively large eyes and approximated orbits in particular. The unique interorbital condition in Saimiri is an example of the effects of life history on skeletal morphology. © 1995 Wiley-Liss, Inc.     

A new lizard-like reptile (Diapsida: Lepidosauromorpha) from the Middle Jurassic of England

The Middle Jurassic (Late Bathonian) Mammal Bed at Kirtlington, Oxfordshire, has yielded a rich assemblage of small vertebrates including mammals, frogs, salamanders and small reptiles. This paper describes the skull of a new diapsid reptile, Marmorelta oxoniensis, which was common within the fauna. The skull is gracile with large orbits and relatively long external nares. The frontals and parietals are single in the adult and almost separated on the roof of the skull by the postfrontals. The temporal region shows a mosaic of primitive and derived character states. The quadrate was firmly held to the skull by a large squamosal and a small, free, quadratojugal was retained. However, reduction of the anterior process of the quadratojugal and of the posterior process of the jugal left the lower temporal fenestra broadly open. Comparison with other diapsids leads to the conclusion that Marmoretta was a lepidosauromorph, most probably the sister taxon of Lepidosauria.     

纤细滇美龙(Dianmeisaurus gracilis Shang & Li,2015)新材料

根据一件产自云南罗平中三叠世关岭组Ⅱ段的新标本并结合产自相同地点和地层中的模式标本对纤细滇美龙(Dianmeisaurus gracilis Shang & Li,2015)进行了详细研究.原模式标本暴露其腹而,而新标本暴露其背面,两者互相补充提供了更完整、精确的纤细滇美龙解剖学信息.新材料揭示该种具有非常短小的吻部,眶前区的长度不仅短于眶后区长度,甚至短于眼眶的长度;外鼻孔小且位置靠前,即鼻孔前区的长度短于鼻孔后缘与眼眶前缘之间的距离;由两额骨构成的眼眶间隔非常狭窄,宽度小于顶骨平台宽度的1/3;额骨前后两端均具渐尖的突起;顶骨后部不收缩,顶骨平台后缘呈深V型.补充了新信息和包含更多属种(如Dawazisaurus)后的系统发育学分析支持了之前滇美龙和滇东龙互为姊妹群的结论,同时它们和马家山龙、滇肿龙、贵州龙和大洼子龙一起构成了一个仅由中国的属种组成的单系类群.与欧洲肿肋龙类群(Dactylosaurus,Anarosaurus,Serpianosaurus和Neusticosaurus)相比,这一单系类群与幻龙类有更近的亲缘关系.     

The cranial anatomy,ontogeny, and relationships of Karpinskiosaurus secundus (Amalitzky) (Seymouriamorpha,Karpinskiosauridae) from the Upper Permian of European Russia

The Upper Permian seymouriamorph tetrapod Karpinskiosaurus from European Russia includes two species: Karpinskiosaurus secundus and Karpinskiosaurus ultimus. Karpinskiosaurus secundus is represented by two specimens with skull lengths of about 75 mm. All specimens of K. ultimus are smaller than those of K. secundus. Revision of the cranial anatomy of all previously known and several new specimens of Karpinskiosaurus shows that the specimens of K. secundus and most of the specimens of K. ultimus represent the ontogenetic series of one species: K. secundus. The holotype specimen of K. ultimus requires revision, with the aim to find out whether it represents a second species of Karpinskiosaurus or not. The available material permits new reconstructions of the largest, holotype skull, and one smaller skull with a length of about 36 mm. Karpinskiosaurus secundus is included in a cladistic analysis for the first time here. The analysis shows it to form a sister taxon to Discosauriscidae. The clade comprising Karpinskiosaurus secundus plus Discosauriscidae forms a sister group to Seymouriidae. Karpinskiosaurus secundus has a large postorbital and a short preorbital region, and the orbits are placed in the posterior portion of the anterior half of the skull length. Among all seymouriamorphs, such cranial proportions are exhibited only by the largest known specimens of Discosauriscus austriacus. None of the specimens of K. secundus described here exhibits the presence of sensory grooves; thus, all specimens composing the ontogenetic sequence of K. secundus are considered to be terrestrial. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010.     

Cranial anatomy of Ignacius graybullianus and the affinities of the Plesiadapiformes

A nearly complete cranium of Ignacius graybullianus provides increased understanding of the cranial anatomy of Plesiadapiformes. In nearly all details of cranial anatomy, Ignacius differs markedly from primates. USNM 421608 exhibits a long tapering snout, small widely spaced orbits, and a complete lack of postorbital process or bar. Large olfactory bulbs are inferred from the wide interorbital space. The marked flare of the zygomatic arches suggests that Ignacius possessed large and powerful temporal muscles. The basicranial region is particularly well preserved and reveals a distinct suture between the petrosal bone and an entotympanic bulla. This suture is visible on both the left and right sides of the skull and dispels the hypothesis that Ignacius and, by inference, other Plesiadapiformes share the primate synapomorphy of a petrosal bulla. To test the phylogenetic position of Ignacius, cranial characters were identified and scored for Ignacius, Plesiadapis, Cynocephalus, and a number of primates, bats, and scandentians. Two erinaceomorph insectivores were also included to allow the assessment of archontan monophyly. These characters were incorporated into a maximum-parsimony analysis to determine the phylogenetic position of Plesiadapiformes. There are several important phylogenetic conclusions that can be inferred from this analysis: 1) Ignacius and Plesiadapis make up a monophyletic clade; 2) Plesiadapiformes may be the sister group of Dermoptera; 3) Scandentia, not Plesiadapiformes, is the sister group of Primates; and 4) Primates, plesiadapiforms, bats, colugos, and scandentians may not form a monophyletic clade Archonta. Consequently, the taxon Archonta is in need of review. © 1992 Wiley-Liss, Inc.     

Blood group genetic variations in inhabitants of West New Guinea, with a map of the villages and linguistic groups of South West New Guinea

In non-primates the nasal fossa manifests a posterior intrasphenoidal extension, accommodating the hindermost ethmoturbinals and an accompanying series of paranasal ectoturbinal cavities within the interorbital region. Primitive primates (e.g. Tarsius, Hapale, Saimiri) show phylogenetic loss of these cavities (together with the interorbital region and posterior ethmoturbinals) and the presence of an interorbital septum: a maxillary sinus only is present and the frontal and sphenoid bones are not pneumatized. In some phylogenetically advanced primates (e.g. Lagothrix, Alouatta) trends of cerebral enlargement and resultant separation of the orbits have induced some degree of reappearance of the lost interorbital territory, and an enhanced but variable pattern of peranasal pneumatization, involving the frontal and the sphenoid. Details of this pattern are described for Tarsius, Saimiri, Hapale, Cebus, Lagothrix and Alouatta, distinction being made between homologous and merely analogous paranasal cavities.     

CRANIAL CREST DEVELOPMENT IN THE AZHDARCHOID PTEROSAUR TUPUXUARA, WITH A REVIEW OF THE GENUS AND TAPEJARID MONOPHYLY

Abstract: A portion of pterosaur skull from the Romualdo Member of the Santana Formation (?Albian–?Turonian, Cretaceous) of north‐east Brazil provides new data on the morphology and ontogeny of azhdarchoid pterosaur cranial crests. The specimen consists of parts of the cranial bones posterodorsal to the nasoantorbital fenestra, including partial nasals, lacrimals, frontals and possibly the parietals. A posterodorsally directed premaxillary crest with a concave posterior border is located dorsal to the posterior border of the nasoantorbital fenestra. A well‐defined suture indicates overlapping, posterodorsally directed growth of the premaxilla over the skull roof, suggesting that the generation of the premaxillary crest is a late ontogenetic feature and thus probably related to sexual display. The systematics of Tupuxuara and its relationship to other azhdarchoids is reviewed and a cladistic analysis of the group is presented. Tupuxuara is found to be the sister‐taxon to Azhdarchidae. Tupuxuara longicristatus Kellner and Campos, 1988 is argued to be the only valid named species in this genus and Thalassodromeus Kellner and Campos, 2002 is considered a junior subjective synonym of this taxon. As originally conceived, Tapejaridae is paraphyletic: a new, more restrictive version of Tapejaridae (including Tapejara and Sinopterus dongi) might exist, but its monophyly is weakly supported. Furthermore, Tapejara was found to be paraphyletic in all trees.     

A SKULL OF THE GIANT BONY‐TOOTHED BIRD DASORNIS (AVES: PELAGORNITHIDAE) FROM THE LOWER EOCENE OF THE ISLE OF SHEPPEY

Abstract: The first substantial skull of a very large Paleogene bony‐toothed bird (Pelagornithidae) is described from the Lower Eocene London Clay of the Isle of Sheppey in England. The specimen is assigned to Dasornis emuinus (Bowerbank), based on a taxonomic revision of the large London Clay Pelagornithidae. Very large bony‐toothed birds from the London Clay were known previously from fragmentary remains of non‐comparable skeletal elements only, and Dasornis londinensis Owen, Argillornis emuinus (Bowerbank), A. longipennis Owen, and Neptuniavis miranda Harrison and Walker are considered junior synonyms of D. emuinus. The new specimen allows a definitive assignment of Dasornis to the Pelagornithidae and documents that this taxon closely resembles other bony‐toothed birds in cranial morphology. It is hypothesized that giant size (i.e. a wingspan above 4 m) evolved only once within Pelagornithidae and that Dasornis emuinus is the sister taxon of the giant Neogene bony‐toothed birds, which share a derived wing morphology.     

Late Oligocene mousebird converges on parrots in skull morphology

A new fossil stem group representative of Coliiformes (mousebirds) with a remarkable skull morphology is described from the late Oligocene of Germany. Oligocolius psittacocephalon sp. nov. for the first time preserves the skull of a post‐Eocene fossil mousebird. This exhibits a combination of skull features unknown from any other bird and converges on the skull of parrots in that the beak is separated from the cranium by a marked nasofrontal hinge and in that the interorbital part of the frontal bones is very wide. In addition, the mandible of the new species exhibits long retroarticular processes, which are unexpected because unlike in other coliiform birds exhibiting this feature, the short beak was probably not used for probing in substrate. It is hypothesized that the retroarticular processes of O. psittacocephalon instead served for a particular wide and forceful opening of the beak. Eight large fruit stones are situated in the area of the digestive tract of the new species. Preservation of most of these in a well‐delimited cluster in the region of the upper oesophagus suggests that, unlike in modern mousebirds, O. psittacocephalon had a crop. The new fossil shows that late Oligocene European stem group Coliiformes significantly differed from their extant relatives in morphology and probably also in feeding ecology.     

A new stereospondyl from the German Middle Triassic, and the origin of the Metoposauridae

Recent finds of well-preserved temnospondyl skeletons from the Lower Keuper (Ladinian, Middle Triassic) in southern Germany are assigned to a new genus and species, Callistomordax kugleri . This taxon is characterized by the following autapomorphies: (1) wide unpaired frontal; (2) vomerine fangs greatly enlarged to occupy entire width of element; (3) intercentra elongated and massive, anterior face being convex; (4) humerus semilunar with enlarged deltopectoral crest; (5) cleithrum strongly curved and bow-shaped; (6) trunk extremely elongated to reach three times the length of the skull. Callistomordax shares with the Metoposauridae the pattern of dermal ornamentation, the proportion of both posterior skull table and snout, the position of the lacrimal, the morphology of the basicranial region, and the structure of the clavicle and interclavicle. Phylogenetic analysis suggests Callistomordax to be the sister taxon of the Metoposauridae, nested within a grade formed by various trematosaurian taxa. In this assemblage, Lyrocephaliscus and a clade formed by Almasaurus , Rileymillerus , Callistomordax , and the Metoposauridae are sister taxa. In all variants of the cladistic analysis, Callistomordax and the Metoposauridae form immediate sister groups. According to the present findings, neither plagiosaurids nor brachyopoids and rhytidosteids are closely related to this 'trematosaurian' monophylum, although these taxa share a range of homoplasies.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 152 , 79–113.     

Redescription of †Paralycoptera wui Chang & Chou, 1977 (Teleostei: Osteoglossoidei) from the Early Cretaceous of eastern China

?Paralycoptera wui Chang & Chou, 1977 from the Early Cretaceous of China is redescribed herein through a re‐examination of the original materials, as well as observations on some newly collected specimens. The use of the peeling method has revealed much of the new or revised information on its osteology, e.g. aspects of the nasal, infraorbitals, retroarticular, preopercle, extrascapular, basihyal toothplate, epineural, pelvic fin, caudal skeleton and scales. The phylogenetic relationships of ?Paralycoptera and other osteoglossomorphs are re‐evaluated. The cladistic analysis largely agrees with the previous hypothesis that ?Paralycoptera is not a ?lycopterid, but rather a stem osteoglossoid. ?Paralycoptera is excluded from notopteroids and exhibits the following derived characters of the Osteoglossoidei: (1) palatal area behind and below orbit completely covered by infraorbitals; (2) jaw articulation under posterior portion of orbit; (3) opercle depth twice or more its width; (4) first pectoral fin ray much enlarged and long, extending posteriorly beyond origin of pelvic fin. ?Singida and ?Phareodus are regarded as different levels of osteoglossoids above ?Paralycoptera. ?Singida shares the following derived characters with ?Phareodus plus extant osteoglossoids: (1) jaw articulation posterior to orbit; (2) anterior process of hyomandibula in contact with entopterygoid; (3) subopercle small and anterior to opercle. ?Phareodus shares the following derived characters with extant osteoglossoids: (1) supraorbital and otic sensory canals connected; (2) one uroneural; (3) reticulate furrows present over entire scale.     

四川自贡大山铺蜀龙动物群——简报Ⅴ 两栖类

本文对扁头中国短头鲵(新属新种) (Sinobrachyops placenticephalus gen. et sp. nov.) 的形态特征和分类位置进行简述.标本得自著名的恐龙化石产地——自贡大山铺,产出时代为中侏罗世.中国短头鲵是目前迷齿类中在地史上最年轻的一个属.它的发现使迷齿类在地球上生存时代的上限推移到中侏罗世.     

Morphology of the interorbital region of Saimiri sciureus

The skull of the platyrrhine primate Saimiri sciureus is distinguished by a large interorbital fenestra. Juvenile skulls still show a bony interorbital septum with some small gaps. A morphogenetic study was undertaken to better understand the structures of the interorbital region, which represents a linkage between the base of the braincase and the nasal skeleton. Already in early ontogenetic stages a reduction of the posterior portion of the nasal capsule and of the cartilaginous interorbital septum are observed, resulting in the formation of a primary interorbital fenestra. A bony interorbital septum is mainly formed in perinatal age stages by ossification of the presphenoid and by medial fusion of the frontals; the primary interorbital fenestra is retained as a small opening. It only occurs in late juvenile stages when the definitive interorbital fenestra develops by by secondary transformation of bone into a membrane of dense connective tissue; this process is most probably caused by mechanical friction of the very closely approximated eyes of both sides.     

Validation and efficacy of transgenerational mass marking of otoliths in viviparous fish larvae

Transgenerational mass marking of viviparous fish larvae in vivo was validated by intra‐muscular injection of elemental strontium chloride (SrCl₂) in gestating females and detection of the Sr in the otoliths of developing larvae. All otoliths of brown rockfish Sebastes auriculatus larvae produced from SrCl₂‐injected females showed enriched Sr:Ca ratios near the otolith edges, and the signatures did not appear to be affected by the anterior, centre and posterior positions of larvae within the ovary. Results from the present study indicate that transgenerational marking is a highly reliable technique for marking large numbers of extremely small viviparous fish larvae.