Ultrastructure of dental hard tissues of Gondwanatherium and Sudamerica (Mammalia, Gondwanatheria)

Sigogneau-Russell, D., Bonaparte, J. F., Frank. R. M. & Escribano. V. 1991 01 15: Ultrastructure of dental hard tissues of Gondwanatherium and Sudamerica (Mammalia, Gondwanathcria). Lethaia. Vol. 24, pp. 27–38. Oslo. ISSN 0024–1164. The ultrastructural study of small dental fragments of the South American mammals Gondwanarherium Bonaparte, 1986 (late Cretaceous) and Sudamerica Scillato-Yané & Pascual. 1985 (middlc Paleocene). tentatively classified in the Paratheria, his shown the same structural characteristics. In the first genus one notes the presence of a thick, well differentiated, prismatic enamel with prisms of pattcrn I of Boyde and interprismatic substance. In places, lines of apposition are visible. The most internal zone of the enamel is very rich in tubules, which become rarer in the peripheral enamel. The dentinc presents numerous parallel tubules, sometimes continuing into the enamel. The presence of enamel over the whole height of the tooth fragment indicates that the hypsodonty here would be due almost entirely to lengthening of the crown. While structurally resembling the specimen of Gondwunarherium, the dental fragment of Sudamerica differs in its enamel locally constituted of open prisms. In spite of the further differences in lobation and enamel folding, the similar kind of hypsodonty met with in both genera studied tends to support their grouping in the order Gondwanatheria Moncs, 1987. an order which may well be situated outside the Tribosphenida. On the contrary, the abscnce. in thc fragments under study. of the dental specializations seen in typical Xenarthra, such as enamel rcduction. suggest that the morphologic resemblance observed between these genera of the Gondwanatheria and Xenarthra might be a phenomenon of convergence rather than of phyletic relationship. However. diversity within Gondwanathcria, and our poor knowledge of the carliest stages of the complex superorder Xenarthra, do not permit at the moment any definitive statement as to their phylogenetic relationships. Teeth, ultrastructure, Gondwanarheria. Xenarthra.     

New hominid fossils from Woranso‐Mille (Central Afar,Ethiopia) and taxonomy of early Australopithecus

The phylogenetic relationship between Australopithecus anamensis and Australopithecus afarensis has been hypothesized as ancestor‐descendant. However, the weakest part of this hypothesis has been the absence of fossil samples between 3.6 and 3.9 million years ago. Here we describe new fossil specimens from the Woranso‐Mille site in Ethiopia that are directly relevant to this issue. They derive from sediments chronometrically dated to 3.57–3.8 million years ago. The new fossil specimens are largely isolated teeth, partial mandibles, and maxillae, and some postcranial fragments. However, they shed some light on the relationships between Au. anamensis and Au. afarensis. The dental morphology shows closer affinity with Au. anamensis from Allia Bay/Kanapoi (Kenya) and Asa Issie (Ethiopia) than with Au. afarensis from Hadar (Ethiopia). However, they are intermediate in dental and mandibular morphology between Au. anamensis and the older Au. afarensis material from Laetoli. The new fossils lend strong support to the hypothesized ancestor‐descendant relationship between these two early Australopithecus species. The Woranso‐Mille hominids cannot be unequivocally assigned to either taxon due to their dental morphological intermediacy. This could be an indication that the Kanapoi, Allia Bay, and Asa Issie Au. anamensis is the primitive form of Au. afarensis at Hadar with the Laetoli and Woranso‐Mille populations sampling a mosaic of morphological features from both ends. It is particularly difficult to draw a line between Au. anamensis and Au. afarensis in light of the new discoveries from Woranso‐Mille. The morphology provides no evidence that Au. afarensis and Au. anamensis represent distinct taxa. Am J Phys Anthropol 2010. © 2009 Wiley‐Liss, Inc.     

The Phylogenetic Affinities of the Enigmatic Mammalian Clade Gondwanatheria

Gondwanatheria is a group of extinct mammals known from the Cretaceous and Paleogene of Gondwana. Resolution of the phylogenetic affinities of gondwanatherians has proven problematical, with the group currently considered Mammalia incertae sedis. We briefly review the morphology of known gondwanatherians, and argue that isolated upper premolars and a partial dentary preserving a blade-like p4 originally referred to the ferugliotheriid gondwanatherian Ferugliotherium windhauseni but subsequently identified as Multituberculata incertae sedis do indeed belong to F. windhauseni. We also suggest that the recently described ?cimolodontan multituberculate Argentodites coloniensis, based on an isolated lower premolar, may in fact be an unworn p4 of Ferugliotherium or a closely related taxon. We present the first phylogenetic analyses to include gondwanatherians, using maximum parsimony and Bayesian methods. Both methods place Ferugliotherium and sudamericid gondwanatherians in a clade with cimolodontan and “plagiaulacidan” multituberculates, although relationships within this clade are largely unresolved. The Gondwanatheria + Multituberculata clade supported here may reflect the convergent evolution of similar dental features, but it is the best supported hypothesis based on currently available data. However, denser sampling of multituberculate taxa and the discovery of more complete gondwanatherian fossils will be required to clarify the precise relationship between gondwanatherians and multituberculates, specifically to determine whether or not gondwanatherians are members of Multituberculata. We hypothesize that the anterior molariforms of sudamericid gondwanatherians evolved from blade-like precursors similar to the p4 of Ferugliotherium, possibly in response to the appearance of grasses in Gondwana during the Cretaceous.     

Taxonomic affinities of the enigmatic Prionogale breviceps,early Miocene,Kenya

Prionogale breviceps is a tiny carnivorous mammal from the early Miocene of eastern Africa. Originally, specimens were interpreted as the adult morphology of the taxon. The dentition did not obviously align Prionogale with the carnivorous lineages present in Afro-Arabia during the early Miocene: Hyaenodonta and Carnivora. When Namasector was discovered in Namibia, the small taxa were placed together in Prionogalidae and aligned with Hyaenodonta. In this study, based on comparisons to hyaenodont specimens preserving deciduous dentition, the holotype of Prionogale is reinterpreted as preserving dP3 and dP4. Some of the lower dental specimens attributed to the taxon preserve dp4. The holotype of Namasector also preserves deciduous dental material. A phylogenetic analysis that includes deciduous dental characters for a broader sample of hyaenodonts resolved Prionogalidae as a clade. Understanding of the deciduous dentition of Prionogale allows future analyses to compare homologous morphology, and to explore the environmental factors that shaped carnivorous mammal evolution through the Miocene.     

New proconsuloid postcranials from the early Miocene of Kenya

New early Miocene forelimb fossils have been recovered from the Songhor and Lower Kapurtay localities in southwestern Kenya. We describe four specimens that are similar in size and functional capabilities. Their specific allocation is problematic but these forelimb specimens must belong to either Rangwapithecus gordoni or Proconsul africanus. If these new postcranial specimens should belong to R. gordoni, on the basis of size and common dental specimens found at Songhor, they represent a new elbow complex. The morphology of these fossils is anatomically and functionally similar to that of Proconsul. The proconsuloid elbow complex allows extensive forelimb rotations and is capable of performing arboreal quadrupedalism and climbing activities. No suspensory adaptations are apparent. The proconsuloid elbow complex remains a good ancestral condition for hominoid primates.     

ANATOMY AND RELATIONSHIPS OF SOME DEVONIAN PROGYMNOSPERMS FROM NEW YORK

New specimens of Triloboxylon ashlandicum Matten and Banks (1966) show additional features of the anatomy and morphology of the plant. The primary xylem has strands of parenchyma associated with trace formation. Secondary xylem and phloem are described for the first time. The outer cortex, previously unknown, contains hypodermal fiber strands. The ultimate appendages are much divided and planated. Specimens of Aneurophyton hallii (Arnold, 1940) were reexamined and newly prepared. They are transferred to the genus Triloboxylon as T. hallii. Triloboxylon is transferred from Pteropsida-Incertae Sedis to the Aneurophytales. New information on the morphology and anatomy of Telraxylopteris is presented and its identity to Sphenoxylon is supported.     

Additional mandibles of Rangwapithecus gordoni,an early Miocene catarrhine from the Tinderet localities of Western Kenya

Two catarrhine mandibles and five isolated teeth have been discovered from Early Miocene localities in Western Kenya. One mandible comes from the well‐known locality of Songhor whereas the other is from a newly discovered locality, Lower Kapurtay, located near Songhor. The mandibles both can clearly be assigned to the species Rangwapithecus gordoni based on molar morphology, which is unique among Early Miocene catarrhines. The isolated specimens can be assigned to Rangwapithecus based on their similarities in morphology to the homologues preserved in the two mandibles. These specimens provide important new information about the dentognathic morphology of Rangwapithecus, which is described in detail. The mandible from Songhor (KNM‐SO 22228) represents the first definitive female mandible of Rangwapithecus. The Lower Kapurtay mandible (KNM‐KT 31234) appears to be male but is much smaller than another recently described male mandible of this species (KNM‐SO 17500) and the type maxilla (KNM‐SO 700). These specimens enable a reassessment of the attributions of all other mandibles and isolated lower teeth of Rangwapithecus, and we present a complete hypodigm of the mandibular and lower dental material for the species. Finally, we provide some additions to the diagnosis of Rangwapithecus gordoni based on previously unknown morphology. Am J Phys Anthropol 153:341–352, 2014. © 2013 Wiley Periodicals, Inc.     

SYSTEMATICS OF GRATELOUPIA FILICINA (HALYMENIACEAE,RHODOPHYTA), BASED ON rbcL SEQUENCE ANALYSES AND MORPHOLOGICAL EVIDENCE,INCLUDING THE REINSTATEMENT OF G. MINIMA AND THE DESCRIPTION OF G. CAPENSIS SP. NOV.1

Grateloupia filicina (C. Agardh) Lamouroux, originally described from the Mediterranean Sea, has long been considered a textbook example of a marine red alga with a cosmopolitan distribution. An rbcL‐based molecular phylogeny, encompassing samples covering the entire geographic distribution of the species, revealed a plethora of “cryptic” species, whereby the presence of genuine G. filicina is limited to the Mediterranean basin. The phylogeny revealed a strong biogeographic imprint, with specimens from temperate regions resolved in clades composed of species inhabiting the same geographic region. Presence of widely divergent morphologies in the temperate clades indicated that several lineages have converged independently to a G. filicina‐type morphology. Tropical representatives are resolved in a single clade with very uniform G. filicina‐type morphology and pairwise sequence divergences that are lower than the average divergence observed in temperate lineages. This, combined with a lack of clear geographic structure among the tropical lineages, may indicate a more recent divergence with long‐range dispersal capacities. Violations to the biogeographic signal in temperate lineages seemed to be due to either inadequate taxonomy or recent introductions. Grateloupia minima P. & H. Crouan, a taxon placed in synonymy under G. filicina, is reinstated as a separate species distributed in the northeast Atlantic Ocean. Grateloupia capensis sp. nov. is described to accommodate specimens from South Africa with a G. filicina‐type morphology, and G. filicina var. luxurians is elevated to species status. Morphological and anatomical characters were put forward that support the distinctiveness of these three distinct species.     

INVESTIGATIONS OF ANGIOSPERMS FROM THE EOCENE OF NORTH AMERICA: STIPULATE LEAVES OF THE RUBIACEAE INCLUDING A PROBABLE POLYPLOID POPULATION

A comparative study was made of the gross morphology, fine venation and cuticular features of Leitneria fioridana Chapman, the single living representative of the order Leitneriales and Leitneria eocenica (Berry) Brown, presumbaly a related fossil species. In addition to the type material, newly collected fossil specimens were investigated from clay pits in the Middle Eocene, Claiborne Formation, of western Tennessee and Kentucky. Foliate stipules attached to the petioles of several specimens suggest the assignment of this fossil leaf type to the genus Leitneria is incorrect. The nature of the gross morphology, fine venation and cuticular features confirms the misidentification. Previously, various specimens of this fossil leaf type have been placed in eight species of seven genera in seven families of six angiosperm orders, none of which are correct systematically. The gross morphology, venation and cuticular characters of the fossil leaf are distributed among a few extant South American genera of arborescent Rubiaceae. The fossil is an extinct rubiaceous leaf type which cannot be placed within a single modern subfamily, tribe or genus of the family. The organ genus, Paleorubiaceophyllum is proposed for these leaves. Three varieties of a single fossil species, P. eocenicum, are recognized. One variety with epidermal cells nearly twice the size of the others may represent a polyploid population.     

CONCORDANCE OF MOLECULAR AND ULTRASTRUCTURAL DATA IN THE STUDY OF ZOOSPORIC CHLOROCOCCALEAN GREEN ALGAE1

Alternative evolutionary hypotheses generated from features of vegetative cell morphology and motile cell ultra-structure were investigated using a molecular data set. Complete nuclear-encoded small subunit (18S) ribosomal RNA (rRNA) gene sequences were determined for six species (three each) of the chlorococcalean green algae “Neo chloris” and Characium. Based on motile cell ultra-structure, it was previously shown that both genera could be separated into three distinct groups possibly representing three separate orders and two classes of green algae. 18S rRNA gene sequences were also obtained for three additional taxa, Dunaliella parva Lerche, Pediastrum duplex Meyen, and Friedmannia israelensis Chantanachat and Bold. These organisms were selected because each, in turn, is a representative of one of the three ultrastructural groups into which the Neochloris and Characium species are separable. Phylogenetic analyses utilizing the molecular data fully support the ultrastructural findings, suggesting that the similar vegetative cell morphologies observed in these organisms have resulted from convergence.     

Dental anomalies within extant members of the mammalian Order Pilosa

Recent discovery of a supernumerary dental anomaly in two‐toed sloths led to an extensive review of extant sloth specimens to look for additional anomalies. In total, 881 museum specimens were examined. These revealed two primary types of anomalies, hyperdontia (extra teeth) and anodontia (loss of teeth), occurring at a rate of 2.4% (n = 21). Two‐toed sloths, Choloepus, were more likely to have hyperdontia in the anterior dentition, whereas three‐toed sloths, Bradypus, experienced anodontia more frequently with the upper caniniforms. Both genera experienced both anomalies. The majority affected the upper dentition, with only three specimens exhibiting mandibular anomalies. Beyond the patterns of tooth positioning, all anomalies were random with respect to age, sex and geography. A few specimens not counted in the initial assessment expressed incomplete anodontia, indicating that the loss occurred postnatally and was not an embryological anomaly. For Bradypus, the findings provide new support for the hypothesis that the taxon represents a neotenic lineage and opens new possibilities about its relationship to the extinct ground sloths with a suggested rooting above that of the basal position it typically occupies for Folivora.     

STUDIES OF PALEOZOIC FERNS. THE MORPHOLOGY,ANATOMY, AND TAXONOMY OF ANKYROPTERIS GLABRA

Eggert , Donald A. (Yale U., New Haven, Conn.) Studies of Paleozoic ferns. The morphology, anatomy, and taxonomy of Ankyropteris glabra. Amer. Jour. Bot. 46(7): 510–520. Illus. 1959.—This is a morphological and taxonomic study of the American specimens of Ankyropteris, with the exception of A. hendricksi, that previous to this time have been referred to A. grayi, originally a Lower Carboniferous species from England, and to A. glabra, a Middle-Pennsylvanian form from Booneville, Indiana. The problem of the separation of these two closely allied species is discussed, and the original criteria for separation are questioned, on the basis of four specimens described in the present paper. The specimens are from West Mineral, Kans. and Shuler Mine in Iowa. They all have been referred to A. glabra and show extreme plasticity of several features of the plant that is correlated with the size of the stem. The major points of difference between A. glabra and A. grayi are now thought to be the presence of distantly spaced nodes in A. glabra, in contrast to closely spaced ones in A. grayi, and the presence of a small amount of mixed tissue in the stele of A. glabra, compared to a large amount in A. grayi. Well preserved axillary branches were found, whose morphology is typically that of the parent stem. It is suggested that the axillary branches were well developed in the species, and that, therefore, several orders of branching may be present in the American material. Successive orders of branching show diminution and simplification which is interpreted as an expression of the presence of determinate growth in the cauline systems of the plant, an ontogenetic pattern that is not commonly found in living forms.     

MOLECULAR ASSESSMENT OF ULVA SPP. (ULVOPHYCEAE,CHLOROPHYTA) IN THE HAWAIIAN ISLANDS1

Sequences of the nuclear internal transcribed spacer 1 (ITS1) region and the chloroplast rbcL gene were obtained from 86 specimens of Ulva (including “Enteromorpha”) from five of the main Hawaiian Islands. These 86 specimens were divided into 11 operational taxonomic units (OTUs) based on analyses of primary sequence data and comparisons of ITS1 secondary structure. Of the 11 OTUs, six have not previously been reported from anywhere in the world. Only three represented exact sequence matches to named species (Ulva lactuca L., syn. U. fasciata Delile; U. ohnoi Hiraoka et Shimada); two others represented exact sequence matches to unnamed species from Japan and New Zealand. Of the 12 species names currently in use for Hawaiian Ulva, only one, U. lactuca (as U. fasciata), was substantiated. General morphology of the specimens did not always correspond with molecular OTUs; for example, reticulate thallus morphology, previously considered diagnostic for the species U. reticulata Forssk., was expressed in thalli assigned to U. ohnoi and to one of the novel OTUs. This finding confirms a number of recent studies and provides further support for a molecular species concept for Ulva. These results suggest that Ulva populations in tropical and subtropical regions consist of species that are largely unique to these areas, for which the application of names based on types from temperate and boreal European and North American waters is inappropriate. Ulva ohnoi, a “green tide” species, is reported from Hawaii for the first time.     

Dental agenesis as evidence of possible genetic isolation in the colobine monkey,Rhinopithecus roxellana

In a sample of 24 skulls of the rare colobine monkey of China,Rhinopithecus roxellana, agenesis of the permanent upper third premolar was observed in 8 specimens (33%). Two of the individuals (8% of the sample) showed unilateral agenesis, while six (25%) showed bilateral agenesis. All of the affected individuals appear to have originated from two areas in central Sichuan Province in China, Moupin (Baoxing) and Wen Chuan, that lie approximately 120 km apart on the western rim of the Sichuan Basin. In this species agenesis of the upper third premolar was generally accompanied by rotation of the upper fourth premolar, but not by any other observable variation in tooth size, morphology, or number. Further, dental agenesis in this species appears to have had no effect craniofacial morphology. In certain human populations, a high prevalence of dental agenesis has been associated with small population effects and genetic isolation. Premolar agenesis inR. roxellana can probably be traced to a similar origin. Reconstruction of the evolutionary history of the species indicates that a severe reduction of the geographic range of the species occurred as a result of climatic deterioration during the Late Pleistocene. Although large numbers of populations appear to have become extinct, others survived to give rise to the modern populations of the species that inhabit China today. The two populations showing a high prevalence of premolar agenesis appear to have originated from one that passed through a population bottleneck and suffered the consequences of founder effect in the Late Pleistocene. This interpretation is supported by evidence of dental agenesis in a population of an insectivore species from the same region and by the fact that premolar agenesis is not found in any of the other species ofRhinopithecus. There is no evidence to support the interpretation that dental agenesis inR. roxellana is due to natural selection, mutation or an evolutionary force other than a small population effect.     

STRUCTURE AND AFFINITIES OF PROTOSTIGMARIA EGGERTIANA

Investigation of new, more or less complete specimens of Protostigmaria eggertiana reveals that it is a large, multilobed rooting structure produced at the base of a small arborescent lycopod. The trunk bears scars marking the point of attachment of leaves, and can be compared to the stem of Lepidodendropsis. In the largest specimens the root-bearing base is divided into about 13 lobes. In smaller, presumably younger specimens, fewer lobes are present. The roots are arranged on the lobes in series and orthostichies comparable to those of Isoetes. New observations on five- and six-lobed Isoetes plants reveal a closer correspondence between Protostigmaria and Isoetes than previously recognized; and support suspected homologies among many fossil and extant lycopsid rooting structures.     

Molar crown inner structural organization in Javanese Homo erectus

This contribution investigates the inner organizational pattern (tooth tissue proportions and enamel–dentine junction morphology) of seven Homo erectus permanent molar crowns from the late Lower‐early Middle Pleistocene Kabuh Formation of the Sangiran Dome (Central Java, Indonesia). The previous study of their external characteristics confirmed the degree of time‐related structural reduction occurred in Javanese H. erectus, and also revealed a combination of nonmetric features which are rare in the Lower and early Middle Pleistocene dental record, but more frequently found in recent humans. In accordance with their outer occlusal morphology, the specimens exhibit a set of derived internal features, such as thick to hyperthick enamel, an incomplete expression of the crest patterns at the enamel–dentine junction (EDJ) level, a sharp EDJ topography. As a whole, these features differ from those expressed in some penecontemporaneous specimens/samples representing African H. erectus/ergaster and H. heidelbergensis, as well as in Neanderthals, but occur in recent human populations. Further research in virtual dental paleoanthropology to be developed at macroregional scale would clarify the polarity and intensity of the intermittent exchanges between continental and insular Southeast Asia around the Lower to Middle Pleistocene boundary, as well as should shed light on the still poorly understood longitudinal evolutionary dynamics across continental Asia. Am J Phys Anthropol 156:148–157, 2015 © 2014 Wiley Periodicals, Inc.     

STEMS AND LEAVES OF CUNNINGHAMIOSTROBUS GOEDERTII FROM THE OLIGOCENE OF WASHINGTON

This report is based on nine specimens of fossil conifer stems and leaves from the Early Oligocene Jansen Creek Member of the Makah Formation. They were collected along the northern shore of the Olympic Peninsula of Washington. The fossils are preserved as siliceous permineralizations and were exposed in surface view along rock fractures. Details of leaf morphology and epidermal construction appear on fracture surfaces of certain specimens while the cellular construction of the leaves and twigs is visible in thin sections. Leaves are dorsiventrally flattened, attached to twigs that contain up to four growth increments of secondary xylem, up to 2.3 cm long, 3.5 mm wide, and have parallel margins with minute teeth. The leaves are about 0.5 mm thick and have a central vascular strand surrounded by transfusion tissue. A large resin canal occurs abaxial to the vascular strand, and generally two additional resin canals occur in the mesophyll near each leaf margin. Leaves are mostly hypostomatic, with sunken stomata in two longitudinal bands, one to each side of the midline of the leaf and each containing eight to 13 longitudinal rows of stomata. Several unusual anatomical features in the stems also occur in the peduncle and cone axis of seed cones described as Cunninghamiostrobus goedertii, which occurs at the same locality. Thus, the leafy twigs belong to the same species as produced the cones. The cones, leaves, and shoots of this Tertiary conifer are similar to those of modern Cunninghamia but differ from the living species in several respects.     

The Skull of Stephanomys and a Review of Malpaisomys Relationships (Rodentia: Muridae): Taxonomic Incongruence in Murids

The cranial morphology of the extinct murid genus Stephanomys, previously known only by dental remains, is described here on the basis of partial skulls of three species of Pliocene age. Important cranial characters of the genus are a robust rostrum, a high zygoma, a wide zygomatic arch, a narrow interorbit, a large orbit, and an optic foramen in the backward position. In addition to some dental characters, Stephanomys shares most of these cranial traits with the extinct Malpaisomys from the Canary Islands. Some of these traits may be linked to the development of large eyes and life in a rocky environment. The peculiar dental pattern of Stephanomys (stephanodonty) is also present in some recent murids (Oenomys and Thamnomys) having a different skull morphology. A comparison with nine other extant genera of murids verified the relationship among Malpaisomys, Stephanomys, and Acomys, supporting our previous conclusion. Phenetic and cladistic analyses of 17 cranial and 23 dental characters show that skull morphology is phylogenetically informative but highly convergent and incongruent with other partial evidence based on dental and biochemical characters. The combined analyses of skull and teeth illustrate a case of mosaic evolution in murids.     

Australopithecus afarensis and the single species hypothesis

Ferguson (1989) has recently argued that the variability seen in the fossils assigned toA. afarensis is far more than expected for a single hominid species, and therefore proposes they represent multiple taxa. In particular, he utilizes data on variation in dental metrics and in premolar morphology in support of this hypothesis. A re-evaluation of these data finds the above conclusion to be unwarranted. Variation in dental metrics providesno basis for separating this sample into multiple taxa, regardless of the analog that is used (i.e. modern primate species or fossil hominid species). Additionally, data on P₃ morphology indicate that thepattern of variation seen in the Laetoli/Hadar sample is comparable to the sexual variation seenwithin a single hominoid species. Overall, the balance of the evidence at present indicates that the fossils from Laetoli and Hadar represent a single hominid species,A. afarensis.     

Feeding mechanisms of the sauropod dinosaurs Brachiosaurus,Camarasaurus, Diplodocus and Dicraeosaurus

Skull morphologies and dental wear patterns have been examined in four sauropod genera to evaluate their probable feeding mechanisms. Wear facets on teeth are generally confined to their apices in Brachiosaurus and Dicraeosaurus and they are sometimes also present on the mesial and distal carinae. Skull morphology and dental wear patterns in Diplodocus and Dicraeosaurus are consistent with a raking motion of the jaws during feeding. Diplodocus became mechanically adapted to feed in this way by evolving anteriorly directed teeth in the premaxilla and mesial parts of the maxilla, and by changing the direction of jaw adduction relative to the long axis of the skull. Similar features are present in the few known skulls of Apatosaurus and they may also have been present in Dicraeosaurus. In Brachiosaurus dental wear patterns also imply a raking motion of the jaws, although the more robust skull and teeth and the more vertically directed action of the jaw adductor muscles have led some to suggest the possibility of isognathous occlusion. Camarasaurus employed a powerful bite in its feeding, possibly with slight propaliny of the lower jaw, and its skull was modified to cope with increased stresses arising from mastication. Archaic sauropods appear largely to have employed isognathic occlusion in chopping off vegetation. The raking motion employed by diplodocids and dicraeosaurids was an advanced mode of cropping and stripping, linked evolutionarily to their highly apomorphic cranial morphology.