A new burnetiamorph (Therapsida: Biarmosuchia) from the Lower Beaufort Group of South Africa

The basal clade Burnetiamorpha is known from only two specimens representing two genera, Proburnetia from the Severodvinskian horizon of the Vyatka River Basin in the Kotelnich district of Russia, and Burnetia from the Dicynodon Assemblage Zone of the Beaufort Group of South Africa. Both genera are of Late Tatarian (Late Permian) age. This paper describes the cranial morphology of a new genus of burnetiamorph, Bullacephalus , from the Late Permian Tapinocephalus Assemblage Zone of the Beaufort Group of South Africa. It is known from a relatively complete skull and lower jaw and is the best preserved burnetiamorph yet discovered. Apart from being the oldest member of the clade, Bullacephalus is also morphologically the least derived and provides new evidence on the phylogeny of this poorly understood group of basal therapsids.     

A new basal dicynodont from the Upper Permian of South Africa

The skull of a small anomodont therapsid, from the Tapinocephalus Assemblage Zone (Abrahamskraal Formation, Beaufort Group, Upper Permian) in Northern Cape Province, South Africa, represents a new basal dicynodont and is described in detail. Colobodectes cluveri gen. et sp. nov. is distinguished from other dicynodonts by an anteroposteriorly extensive caniniform process, parietals that were broadly overlapped posterolaterally by posterodorsal processes of the postorbitals, diverging anterior palatal ridges, and a dorsoventrally low foramen magnum. A phylogenetic analysis indicates that Colobodectes is the basalmost member of a dicynodont clade that excludes Eodicynodon . This position is not particularly strong, as two additional steps are needed to make Colobodectes and Eodicynodon oosthuizeni exchange places on the most parsimonious tree. Another discovery of the phylogenetic analysis is that there is little basis for recognizing Eodicynodon oelofseni as the closest relative of E. oosthuizeni . The former species is identified as the sister taxon of a clade that includes the latter and all other dicynodonts.     

Cranial structure and relationships of the Liassic mammal Sinoconodon*

The skull of the 'Rhaeto-Liassic' mammal Sinoconodon changchiawaensis (Young) from the lower Lufeng Series of China is described. It is characterized by a relatively larger and more robust dentary condyle and a greater reduction of the post-dentary bones than are present in the Morganucodontidae, Kuehneotheriidae, and Dinnetherium. In other aspects Sinoconodon is more primitive; precise post-canine occlusion is lacking, the mandibular symphysis is deep, the jaw articulation lies below a line projected through the apices of the teeth, the pterygoparoccipital foramen is large and the post-canine teeth cannot be divided into molars and premolars. The jaw articulation and braincase of Sinoconodon are compared with those of the two cynodont therapsids Probainognathus and Thrinaxodon. It is concluded that in the transition from therapsid to mammal the medial surface of the groove in the squamosal housing the quadrate was lost and, as a result, in Sinoconodon, Morganucodon and Dinnetherium the hollow medial surface of the quadrate abutted directly against the paroccipital process. A definition for the Class Mammalia is given. It is suggested that the three-boned middle ear was present in Cretaceous triconodonts, and that it probably arose independently in the lines leading to multituberculates and Cretaceous therians. The structure of recently discovered 'Rhaeto-Liassic' mammals and that of Sinoconodon indicates that there was greater diversity among the earliest known mammals than was previously thought.     

Cenomanian Cephalopods of Bellezma-Aures mountains,NE Algeria: Taxonomy and biostratigraphy

The Cenomanian Smail Marl Formation in the Bellezma and Aures mountains of northeastern Algeria is well exposed and rich in macroinvertebrates, including cephalopods. Twenty-five ammonite species and two nautiloid species from the Thénièt El Manchar, Djebel Metrassi and Djebel Bouarif sections are described and seven assemblage zones proposed as provisional working units: Mantelliceras mantelli–Sharpeiceras laticlavium Assemblage Zone (lower Cenomanian), Mantelliceras dixoni Assemblage Zone (uppermost lower Cenomanian), Cunningtoniceras inerme Assemblage Zone (lower middle Cenomanian), Acanthoceras rhotomagense and Acanthoceras cf. rhotomagense Assemblage Zone (middle middle Cenomanian), Acanthoceras amphibolum Assemblage Zone (upper middle Cenomanian), Acanthoceras jukesbrownei Assemblage Zone (uppermost middle Cenomanian), and Neolobites vibrayeanus Assemblage Zone (lower upper Cenomanian). The lower Cenomanian Mantelliceras mantelli–Sharpeiceras laticlavium Zone is subdivided into the Submantelliceras aumalense, Sharpeiceras schlueteri and Mantelliceras saxbii subzones. The base of the middle Cenomanian is marked by the disappearance of species of Mantelliceras and the appearance of the scleractinian coral Aspidiscus cristatus (Lamarck, 1801), associated with Cunningtoniceras inerme (Pervinquière, 1907). The middle Cenomanian contains abundant A. cristatus and species of Acanthoceras, i.e., A. cf. rhotomagense (Defrance in Brongniart, 1822), A. jukesbrownei (Spath, 1926) and A. amphibolum Morrow, 1935. An ammonite zonal scheme is then applied to the Cenomanian exposed in the Bellezma-Aures mountains located on the southern Tethys margin in Algeria.     

Revision of the foraminiferal biozonation scheme in Upper Cretaceous carbonates of the Dezful Embayment,Zagros, Iran: Integrated palaeontological,sedimentological and geochemical investigation

Cretaceous carbonate successions of the Bangestan Group, such as the Sarvak and Ilam formations, are among the most prolific hydrocarbon reserves of the Middle East. However, relatively little is known about their detailed palaeontology and biostratigraphy. Moreover, due to lithological similarity of these carbonate formations recognition of their boundaries in subsurface studies is problematic. To investigate these units, biostratigraphic analyses were carried out on nearly 1100 m of cores, including core plug samples and thin sections prepared from five giant and supergiant oilfields in the northern and southern Dezful Embayment, SW Iran. Accordingly, 59 species of foraminifera (assigned to 43 genera) as well as 11 species of non-foraminifera (10 genera) were recognized. As a result, three biozones were identified, which in stratigraphic order are: Nezzazata-Alveolinids Assemblage Zone; Moncharmontia apenninica-Nezzazatinella-Dicyclina Assemblage Zone; and Rotalia skourensis-algae Assemblage Zone. These are compared with the Wynd's (1965) biozonation scheme, previously introduced in the Zagros area, and a revised scheme is presented. Accordingly, a Cenomanian–Turonian age and a Coniacian–Campanian age are envisaged for the Upper Sarvak and Ilam formations, respectively. In our new biostratigraphic scheme, the Sarvak–Ilam formations boundary is considered to be located above the Moncharmontia apenninica-Nezzazatinella-Dicyclina Assemblage Zone (equivalent of Valvulammina-Dicyclina Assemblage Zone of Wynd, 1965), that is Turonian in age. This zone is bounded by two palaeoexposure surfaces, which correspond approximately to the C–T boundary transitional interval and a post-Turonian, which can be possibly assigned to the Coniacian. Significant sedimentological features of these disconformities include bauxitic–lateritic horizons, karstified profiles and solution-collapsed breccias. Geochemical signatures of these meteorically altered surfaces are also considered to calibrate biofacies and biozones. Finally, we compared our new biozonation scheme with other studies in neighboring areas of SW Iran and the Middle East.     

The Origin of the Mammalian Middle Ear

A functional explanation is presented for the shift of the reptilianarticular and quadrate into the mammalian middle ear to becomethe malleus and incus. Modification of the masticatory apparatusof therapsids results in reduction of stresses on the jaw jointand consequently in reduction of posterior elements of the jaw.In the late therapsid, Bienotherium, the quadrate and post-dentaryjaw bones resemble the mammalian malleus and incus which togetherform a lever. The therapsid articular possesses a downturnedretroarticular process (for insertion of M. depressor mandibulae)homologous with the manubrium (force lever arm) of the malleus.About the time of origin of the mammalian (dentarysquamosal)jaw joint and following the origin of the mammalian depressor,the reptilian depressor is lost. This allows the enlarging reptiliantympanum to become attached to the retroarticular process. Thenew lever system thus formed by articular and quadrate increasesthe sensitivity of the ear and the reptilian one-bone systemis replaced. In early mammals the reflected lamina of the angularmigrates posteriorly with the angle of the dentary so that itcontacts and assumes support of the tympanum. Non-homology ofthe monotreme and therian depressors indicates a multiple originof the mammalian middle ear.     

中国肯氏兽-山西鳄组合带的新发现之三:山西临县的主龙型类

近几十年,华北的二马营组上部地层以产出中国肯氏兽-山西鳄四足动物化石组合而闻名.最近在山西临县白道峪于上覆的铜川组一段发现了中国肯氏兽.本文描述了同一地点同一层位产出的主龙型类化石,包括一具山西鳄的部分骨架以及一些可以归入suchian的主龙类.它们是铜川组一段首次记述的主龙型类.最有鉴定特征的suchian材料包括一个大的髂骨以及一个小的、形状很奇特的、可能是跟骨的骨骼.髂骨可能可以归入一个奇异的波波龙类(poposauroid).因为中国肯氏兽与山西鳄同时出现在白道峪,表明中国肯氏兽-山西鳄组合可以向上延伸到铜川组一段.髂骨与跟骨大小悬殊,可能代表两个从未在中国肯氏兽-山西鳄组合报道过的物种.白道峪发现了特化的波波龙类,支持了波波龙类在中三叠世大量分化的观点.     

Fluctuations in a late devonian-early Mississippian phytoplankton flora of Ohio,U.S.A

Fossil diversity and abundance fluctuations of Late Devonian (Famennian)—Early Mississippian (Tournaisian) phytoplankton assemblages were determined in a stratigraphic sequence of samples in Barberton test core 3, from northeast Ohio. Both diversity and abundance were relatively high during deposition of the lower Chagrin Shale (Upper Devonian) and both decreased during deposition of the upper Chagrin Shale. The overlying Cleveland Shale (Upper Devonian) is characterized by sharply increased abundance with a corresponding (although less marked) increase in diversity; in contrast, both abundance and diversity were at relatively low levels during deposition of the Lower Mississippian Bedford Shale, the uppermost unit of the sequence. These trends agree with the phytoplankton periodicity model presented by Tappan, which indicates large-scale extinction at the end of the Devonian.Many of the acritarch and prasinophycean algal taxa have restricted stratigraphic occurrences within the section studied, making them useful for local and perhaps worldwide correlation. The stratigraphically restricted taxa provide a basis for the proposal of five biostratigraphic zones within the Upper Famennian (Upper Devonian) and a portion of the Lower Mississippian. In ascending order occur: the Gorgonisphaeridium ohioense Assemblage Zone, the Acriora petala Assemblage Zone, the Muraticavea enteichia Range Zone, the Cymatiosphaera labyrinthica Range Zone, and the Navifusa drosera Assemblage Zone.     

Paleobiogeographic implications of the Middle Eocene ostracods from Cairo–Suez district,Egypt

The main targets of this paper are to examine the Middle Eocene ostracod assemblages collected from a succession exposed in the Cairo–Suez district, Egypt and to detect their paleobiogeographical implications. The studied succession is subdivided into two rock units: the Observatory and Qurn formations, in ascending order. The analysis of the ostracod assemblages led to the identification of 18 species, none of which is new. Three local biozones are established, Digmocythere ismaili - Xestoleberis kenawyi Assemblage Zone, Grinioneis moosi - Loxoconcha pseudopunctatella Assemblage Zone and Cativella qurnenis - Soudanella triangulata Assemblage Zone. The multivariate analyses indicate that there are three distinct bioprovinces, one of them represents the North Africa bioprovince, including Egypt, Libya, and Tunisia. The second bioprovince represents the Middle East, including Jordan and Israel. The third represents the West Africa bioprovince, including Senegal, Togo, Ivory Coast and Nigeria. Therefore, this deduction supports a migration of Eocene ostracods along the southern Tethys.     

Kissinella-Christiania Associations in the early Ashgill Foliomena brachiopod fauna of South China

A new ecological unit within the well-documented late Ordovician brachiopod Foliomena fauna is defined on the basis of assemblages from Northern Cuizhou and southern Sichuan. Southwest China. The Kassinella-Chrisriania Association, from the lower Ashgill Linhsiang and Chientsaokou formations, is dominated by the eponymous genera together with species of mainly Dedzetina, Sericoidea and Kozlowskites Foliomena is rare or absent. To date, the Foliomena fauna has been documented only from deep-water environments however, the new association apparently occupied shallower depths, probably in lowermost Benthic Assemblage Zone 3 and uppermost Benthic Assemblage Zone 4, presumably at the shallow end of the depth range of the Foliomere fauna. The incursion of the Foliomena fauna into relatively shallow-water environments may have been encouraged by abnormally low oxygen levels and sparse nutrients together with persistent soft substrates across this part of the Yangtze Platform during the early Ashgill. *** Ordovician, Ashgill, China, brachiopods. F oliomena fauna.     

Feeding mechanism of Epibulus insidiator (Labridae; Teleostei): Evolution of a novel functional system

The feeding mechanism of Epibulus insidiator is unique among fishes, exhibiting the highest degree of jaw protrusion ever described (65% of head length). The functional morphology of the jaw mechanism in Epibulus is analyzed as a case study in the evolution of novel functional systems. The feeding mechanism appears to be driven by unspecialized muscle activity patterns and input forces, that combine with drastically changed bone and ligament morphology to produce extreme jaw protrusion. The primary derived osteological features are the form of the quadrate, interopercle, and elongate premaxilla and lower jaw. Epibulus has a unique vomero-interopercular ligament and enlarged interoperculo-mandibular and premaxilla-maxilla ligaments. The structures of the opercle, maxilla, and much of the neurocranium retain a primitive labrid condition. Many cranial muscles in Epibulus also retain a primitive structural condition, including the levator operculi, expaxialis, sternohyoideus, and adductor mandibulae. The generalized perciform suction feeding pattern of simultaneous peak cranial elevation, gape, and jaw protrusion followed by hyoid depression is retained in Epibulus. Electromyography and high-speed cinematography indicate that patterns of muscle activity during feeding and the kinematic movements of opercular rotation and cranial elevation produce a primitive pattern of force and motion input. Extreme jaw protrusion is produced from this primitive input pattern by several derived kinematic patterns of modified bones and ligaments. The interopercle, quadrate, and maxilla rotate through angles of about 100 degrees, pushing the lower jaw into a protruded position. Analysis of primitive and derived characters at multiple levels of structural and functional organization allows conclusions about the level of design at which change has occurred to produce functional novelties.     

Endothiodont dicynodonts from the Late Permian Kundaram Formation,India

The Kundaram Formation, a Lower Gondwana unit of the Pranhita-Godavari valley, contains the only known Permian dicynodonts in India. This dicynodont assemblage is dominated by Endothiodon. Two species (Endothiodon mahalanobisi sp. nov and Endothiodon uniseries Owen 1879) are described here. This is the first detailed study of Endothiodon from outside Africa. The study shows that Endothiodon jaw articulation permitted propalinal action when the opposing serrations in the upper and the lower jaw teeth occluded with the horn covered areas and helped in shearing and mastication. The presence of oblique wear facets on the upper and lower jaw teeth and a masseter-like muscle suggest a slightly lateral movement of the lower jaw which enhanced slicing of plant matter. A robust stapes with posterolateral notch constitutes the middle ear region though Endothiodon depended mainly on ground vibrations for hearing. Other features included a reduced olfactory sense and large eyes for vigilance and ease of detecting predators. Based on its vertebrate fauna, this fossiliferous Gondwana horizon of the Pranhita-Godavari valley is broadly correlated with the Tropidostoma and/or Cistecephalus Assemblage Zones of the Beaufort Group, Karoo Supergroup, South Africa. It indicates a probable Tatarian age for the Kundaram Formation.     

The trigeminal jaw adductors of primitive snakes and their homologies with the lacertilian jaw adductors

The trigeminal jaw adductor musculature of anilioid snakes is analysed. The group is characterised by primitive characters, viz. the presence of an extensive bodenaponeurosis and of a quadrate aponeurosis. A temporal tendon gives rise to superficial (lb) fibres which are not observed in other snakes: this may be a primitive or a derived feature.
Jaw adductor muscles in snakes are usually subdivided following their relative position in an antero–posterior direction. Lacertilian jaw adductors are subdivided in a transverse plane. A detailed comparison of the anilioid and primitive lacertilian jaw adductors establishes correspondences (homologies) of parts in the transverse plane in both groups. These homologies are corroborated by innervational patterns.
Platynotan lizards are widely accepted as potential snake ancestors. A comparison of homologue jaw adductors shows different evolutionary trends to characterise platynotan lizards and snakes. Theoretically, these findings do not rule out primitive platynotan lizards as snake ancestors. On the basis of the structure of jaw adductors, snakes are to be derived from a primitive lacertilian pattern, be it platynotan or not.     

The Fates of Spiracular Allostoses in Mammals

The human temporal bone includes an upper and a lower allostotic component, which are referred to, respectively, as the pars squamosa and pars tympanica in the adult and as the os squamosum and os tympanicum in the fetus. The consensus is that the former is descended from a cheek bone and the latter from a jaw bone in piscine osleolepipods. However, no corroborating evidence supports this view. It is concluded here that both these allostotic components of the human temporal bone derive from spiracular allostoses, except for the zygomatic process of the pars squamosa, this being a cheek allostosis that has been secondarily united with the squama. This is the first recognition of erstwhile spiracular allostoses in tetrapods. These conclusions challenge the widely accepted Reichert-Gaupp theory as well as the oft-alleged monophyletic status of the mammals and their derivation from therapsid reptiles.     

福建大田崆峒山晚二叠世长兴组的牙形刺

该文记述了福建大田崆峒山晚二叠世长兴组牙形刺化石4属7种,在长兴组上部建立了Neogondo-lella changxingensis-N.deflecta-N.subcarinata组合带,着重分析了崆峒山长兴组牙形刺的分布规律和组合特征。认为该组合可与浙江煤山长兴阶上部牙形刺组合带对比;福建大田崆峒山长兴组仅大致相当于我国华南长兴阶或浙江煤山长兴组的中、上部,其下伏的原属翠屏山组海相碎屑岩应划归长兴阶。     

The lower jaw of an aegialodontid mammal from the Early Cretaceous of Mongolia

A partial lower jaw is described of an aegialodontid mammal, Kielantherium gobiensis Dashzeveg, 1975, from the Guchin Us beds of Mongolia (?Aptian or ?Albian). The jaw has four molars and four or five premolar loci. A count of P5 M4 is argued to be primitive for the Tribosphenida. McKcnna's interpretation of the postcanine dentition of Peramus as P5 M3 is accepted. It follows that the Peramura could not lead to the Tribosphenida, which apparently arose from unknown 'pantotheres' with not less than nine postcanine teeth.     

黔南紫云、晴隆晚石炭世及早二叠世四射珊瑚

通过对黔南地区晚石炭世及早二叠世早期四射珊瑚的研究,建立4个四射珊瑚组合,自下而上：1）Antheria-Pseudotimania组合;2）Nephelophyllum-Eokepingophyllum组合;3）Parawentzellophyllum-Kepingophyllum组合;4）Anfractophyllum组合。组合1、2以Cyathopsidae科分子和Kepingophyllidae科的原始分子为主;组合3以Kepingophyllidae科的典型分子为代表;组合4以Kepingophyllidae的进化型分子为特征。文中描述四射珊瑚9新种及1未定种。     

A New Species of Garjainia Ochev, 1958 (Diapsida: Archosauriformes: Erythrosuchidae) from the Early Triassic of South Africa

A new species of the erythrosuchid archosauriform reptile Garjainia Ochev, 1958 is described on the basis of disarticulated but abundant and well-preserved cranial and postcranial material from the late Early Triassic (late Olenekian) Subzone A of the Cynognathus Assemblage Zone of the Burgersdorp Formation (Beaufort Group) of the Karoo Basin of South Africa. The new species, G. madiba, differs from its unique congener, G. prima from the late Olenekian of European Russia, most notably in having large bony bosses on the lateral surfaces of the jugals and postorbitals. The new species also has more teeth and a proportionately longer postacetabular process of the ilium than G. prima. Analysis of G. madiba bone histology reveals thick compact cortices comprised of highly vascularized, rapidly forming fibro-lamellar bone tissue, similar to Erythrosuchus africanus from Subzone B of the Cynognathus Assemblage Zone. The most notable differences between the two taxa are the predominance of a radiating vascular network and presence of annuli in the limb bones of G. madiba. These features indicate rapid growth rates, consistent with data for many other Triassic archosauriforms, but also a high degree of developmental plasticity as growth remained flexible. The diagnoses of Garjainia and of Erythrosuchidae are addressed and revised. Garjainia madiba is the geologically oldest erythrosuchid known from the Southern Hemisphere, and demonstrates that erythrosuchids achieved a cosmopolitan biogeographical distribution by the end of the Early Triassic, within five million years of the end-Permian mass extinction event. It provides new insights into the diversity of the Subzone A vertebrate assemblage, which partially fills a major gap between classic ‘faunal’ assemblages from the older Lystrosaurus Assemblage Zone (earliest Triassic) and the younger Subzone B of the Cynognathus Assemblage Zone (early Middle Triassic).     

Control of jaw movements in two species of macropodines (Macropus eugenii and Macropus rufus).

The masticatory motor patterns of three tammar wallabies and two red kangaroos were determined by analyzing the pattern of electromyographic (EMG) activity of the jaw adductors and correlating it with lower jaw movements, as recorded by digital video and videoradiography. Transverse jaw movements were limited by the width of the upper incisal arcade. Molars engaged in food breakdown during two distinct occlusal phases characterized by abrupt changes in the direction of working-side hemimandible movement. Separate orthal (Phase I) and transverse (Phase II) trajectories were observed. The working-side lower jaw initially was drawn laterally by the balancing-side medial pterygoid and then orthally by overlapping activity in the balancing- and working-side temporalis and the balancing-side superficial masseter and medial pterygoid. Transverse movement occurred principally via the working-side medial pterygoid and superficial masseter. This pattern contrasted to that of placental herbivores, which are known to break down food when they move the working-side lower jaw transversely along a relatively longer linear path without changing direction during the power stroke. The placental trajectory results from overlapping activity in the working- and balancing-side adductor muscles, suggesting that macropods and placental herbivores have modified the primitive masticatory motor pattern in different ways.