Two new species of Dunkleosteus Lehman, 1956, from the Ohio Shale Formation (USA,Famennian) and the Kettle Point Formation (Canada,Upper Devonian), and a cladistic analysis of the Eubrachythoraci (Placodermi,Arthrodira)

Two pachyosteomorph arthrodires are described from the Upper Devonian of North America, Dunkleosteus raveri sp. nov. and Dunkleosteus amblyodoratus sp. nov. (from the Appalachian and the Michigan basins, respectively). Dunkleosteus raveri sp. nov. is found stratigraphically below the only other species within the Ohio Shale Formation (Famennian), and is characterized by the retention of a primitive dermal ornamentation, and shares with Dunkleosteus terrelli ( Newberry, 1873 ) the possession of articular facets on the parasphenoid. Dunkleosteus amblyodoratus sp. nov. (Kettle Point Formation, Upper Devonian), known only from incomplete nuchal, paranuchal, and parasphenoid plates, also retains dermal tuberculation, and is characterized by tapering articular facets on the parasphenoid. The current study reviews three North American species of Dunkleosteus (Du. terrelli, Du. raveri sp. nov. , and Du. amblyodoratus sp. nov. ). Formerly, two subgroups were recognized within the Dinichthyidae, the terrelli and herzeri groups, which are shown here to be members of distinct clades. Dinichthys herzeri Newberry, 1868 (the namesake member of the herzeri group) is now the sole member of the family Dinichthyidae comb. nov. , and is assigned to the Aspinothoracidi sensu Miles & Dennis, 1979 . Heintzichthys Whitley, 1933 and Gorgonichthys Claypole, 1892 (additional members of the herzeri group) were previously placed among the aspinothoracid arthrodires. Likewise, Holdenius holdeni Dunkle & Bungart, 1942 (the final member of the herzeri group) and Hadrosteus rapax Gross, 1932 are now placed within the Aspinothoracidi. The remaining taxa, formerly assigned to the Dinichthyidae (i.e. Dunkleosteus Lehman, 1956 ; Eastmanosteus Obruchev, 1964 ; Golshanichthys Lelièvre, Janvier, & Goujet, 1981 ; and Westralichthys Long, 1987 ), are united within the Dunkleosteidae Stensiö, 1963 . © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 159 , 195–222.     

湖南晚泥盆世短胸节甲鱼类一新属

本报道的鱼类化石产自湖南石门大京州的上泥盆统写经寺组。在中国泥盆纪地层分区上,化石产地属于华南区的湘、鄂、川交界分区(侯鸿飞等,1988)。该分区泥盆系为一套厚度不超过500m的近岸海相碎屑岩沉积,仅发育中泥盆统上部至上泥盆统(图1),并普遍超覆在志留系温洛克统之上,以角度不整合伏于石炭系或二叠系之下。上泥盆统分两个组,从下而上分别为黄家磴组和写经寺组。黄家磴组系杨敬之、穆恩之(1953)创立,标准地点在湖北长阳马鞍山东端的黄家磴。岩性为砂岩夹少量页岩,植物化石丰富,并含有误订为植物化石的湖北长阳鱼(Changyanophyton hupeiense Sze,1952)。根据植物化石组合面貌,黄家磴组的时代被认为是晚泥盆世早期。写经寺组是谢家荣、刘季辰(1927)创立的。岩性分为二段：下段以泥灰岩为主,含Yunnanella abrupta、Yunnanellina triplicata等腕足类化石;上段以砂页岩为主,含植物化石,其组合面貌除大致与黄家磴组相似外,还有一些新的分子,如Leptophloeum guanzhuangense、Cyclostigma hunanense等。根据动、植物化石群的性质,写经寺组的时代大致与法门期早期相当。多年来,在这一分区的泥盆纪地层里,很少鱼类化石。杨敬之、穆恩之(1953)曾提及在湖北长阳写经寺组底部发现的鱼化石,这些鱼化石与腕足类Yunnanella、Tenticospirifer等共生,但种属并未定出。已记述过的鱼化石只有湖北长阳黄家磴组的长阳鱼(潘江,1962)。     

Vertebrate fossils of the Middle Devonian (Eifelian) Mühlenberg Formation in the Bergisches Land, northwestern Germany

Vertebrate fossils of the lower Middle Devonian Mühlenberg Formation from the western Bergisches Land (eastern Rhenish Massif) are described. The placoderms are represented by disarticulated plates of coccosteid arthrodires and the acanthodians by several ischnacanthid jaw bones and aMachaeracanthus finspine. The arthrodires are assigned to a new species; a new genus has to be erected if remains of trunk and head shield remains are conspecific.     

Distribution and ontogenetic shifts in habitat and abundance of the temperate western blue groper, Achoerodus gouldii (Richardson)

Densities of juveniles western blue groper Achoerodus gouldii on exposed coasts of South Australia were slightly greater on granite than other substrata and increased with increasing bottom relief and declined with depth and increasing exposure. Sub-adults were in higher density on schists than on other substrata, and densities increased with increasing bottom relief, and declined with depth, increasing exposure and increasing algal canopy cover. Adults were in higher densities on schists, and abundances increased with depth and bottom relief. At a mesoscale A. gouldii was absent from gulfs and sheltered bays, except near their entrances, while at offshore islands juveniles were rare, sub-adults were in lower abundance, and adults in greater abundance, compared with mainland coasts. It is hypothesized that sub-adults may swim up to 60 km from settlement locations to exposed areas and to offshore islands. At a geographic scale, abundances of all three size classes of A. gouldii declined significantly from west to east over 2000 km of coast from the centre of the species' range to its eastern edge. Although A. gouldii is a protected species over part of its range, abundance of sub-adults and adults was significantly reduced by fishing, as shown by an index of fishing intensity. The mean size of sub-adult A. gouldii was also correlated with this index, suggesting that the observed geographic trend in sub-adult and adult abundance may be contributed to by fishing. Simulation showed that a high instantaneous fishing mortality rate of 0·8 per annum could explain the maximum observed reductions in mean size of sub-adult A. gouldii .     

31P NMR probes of sipunculan erythrocytes containing the O2-carrying protein hemerythrin

Reported are the first examinations by 31P NMR of erythrocytes containing the non-heme iron O2-carrying protein hemerythrin (Hr). Intact coelomic erythrocytes from the sipunculids Phascolopsis gouldii and Themiste zostericola were shown by 31P NMR to contain O-phosphorylethanolamine and 2-aminoethylphosphonate as the major soluble phosphorus metabolites. This combination of major metabolites appears to be unique to sipunculan erythrocytes. Nucleoside triphosphates and mannose 1-phosphate were present in lower concentrations. The concentration of O-phosphorylethanolamine within P. gouldii erythrocytes was established to be greater than 20 mM. T. zostericola erythrocytes contained relatively high levels of 2-aminoethylphosphonate (on the order of 0.1 M) and lower levels of O-phosphorylethanolamine compared with those of P. gouldii. For P. gouldii and T. zostericola the intracellular pHs were determined to be 7.2 +/- 0.1 and 7.1 +/- 0.1, respectively, in air-equilibrated erythrocytes, and 6.5 +/- 0.1 in anaerobic P. gouldii erythrocytes. O-Phosphorylethanolamine was found to bind weakly to P. gouldii metHr (Kf approximately 7 M-1). This interaction is best characterized by either negative cooperativity or nonspecific binding. O-Phosphorylethanolamine strongly inhibits azide binding to the iron site of P. gouldii metHr at pH 7.2. The rate of azide binding decreases by approximately 85-fold in the presence of 0.33 M O-phosphorylethanolamine. However, neither O-phosphorylethanolamine nor 2-aminoethylphosphonate at 0.33 M was found to have any significant effect on O2 affinity of P. gouldii deoxyHr. Alternative functions for the two metabolites are suggested.     

Some behavioural observations on wild and captive sand monitors, Varanus gouldii (Sauria: Varanidae)

Behavioural observations were made on wild and captive sand monitors in Australia. Threat and defensive postures are described for V. gouldii and reference is made to V. varius and V. spenceri; such behaviour is compared to other species within the family Varanidae. With captive V. gouldii, one case of cannibalism occurred when space was restricted.     

New finds of vertebrates in the Middle Devonian Brandenberg Group (Sauerland, Northwest Germany)

The arthrodires of the Eifelian Brandenberg Group of Hagen-Ambrock and Lasbeck (Northwest Germany) are described. A new tuberculated holonematid arthrodire (Holonema bruehni n. sp.) is represented by plates of the trunk shield and some plates of the head region. The tuberculation is here regarded as the plesiomorphic character of the Holonematidae compared to an ornamentation of ridges in advanced genera. This agrees with the stratigraphical occurrence of the new form. Additionally, several other plates belonging to at least three different taxa of coccosteid arthrodires have been found. Despite the absence of complete skeletons, the placoderm fauna is more diverse than the fauna of the classical Scottish Eifelian localities.     

A second eubrachythoracid arthrodire from Gogo, Western Australia

A new genus of long-snouted, eubrachythoracid arthrodire, Camuropiscis gen. nov., is described from the Upper Devonian Gogo formation of Western Australia. It is represented by two species, the type C. concinnus sp. nov. and C. laidlawi sp. nov. These species are interchangeable with Coccosteus spp. in a sketch cladograin for arthrodires; their phylogenetic relationships are not known in any detail. C. concinnus has a well-preserved pelvic girdle and throws new light on the structure of this element in arthrodires.     

Periodic fluctuations in the pulmonary surfactant system in Gould's wattled bat (Chalinolobus gouldii)

Pulmonary surfactant is a mixture of phospholipids, neutral lipids, and proteins that controls the surface tension of the fluid lining the lung. Surfactant amounts and composition are influenced by such physiological parameters as metabolic rate, activity, body temperature, and ventilation. Microchiropteran bats experience fluctuations in these parameters throughout their natural daily cycle of activity and torpor. The activity cycle of the microchiropteran bat Chalinolobus gouldii was studied over a 24-h period. Bats were maintained in a room at constant ambient temperature (24 degrees C) on an 8L : 16D cycle. Diurnal changes in the amount and composition of surfactant were measured at 4-h intervals throughout a 24-h period. The C. gouldii were most active at 2 a.m. and were torpid at 2 p.m. Alveolar surfactant increased 1.5-fold immediately after arousal. The proportion of disaturated phospholipid remained constant, while surfactant cholesterol levels increased 1.5-fold during torpor. Alveolar cholesterol in C. gouldii was six times lower than in other mammals. Cholesterol appears to function in maintaining surfactant fluidity during torpor in this species of bat.     

A primitive eubrachythoracid arthrodire from Gogo, Western Australia

A new eubrachythoracid arthrodire, Hanytoombsia elegans gen. et sp. nov., is described from the Upper Devonian Gogo formation of Western Australia. Its structure and relationships are discussed within the framework provided by a new sketch cladogram for arthrodires. H. elegans is morphologically similar to Coccosteus cuspidatus , but its relationships among primitive eubrachythoracids are unknown.     

Microsatellite markers for the endemic New Zealand long-tailed bat (Chalinolobus tuberculatus)

The New Zealand long-tailed bat (Chalinolobus tuberculatus) is an endangered species which is now absent from many habitats where they were previously observed. However, population genetics studies have been hampered by the lack of reliable genetic markers. We have developed 13 microsatellite markers, and at least eight of these are suitable for population studies. Nine of these markers also cross-react with the closely related Australian species C. gouldii. These markers may prove of general utility in the genus Chalinolobus.     

A pachyosteomorph arthrodire from Gogo, Western Australia

A new monotypic genus of eubrachythoracid arthrodire is described from die Upper Devonian Gogo Formation of Western Australia as Incisoscutum ritchiei gen. et sp. nov. It is at die pachyosteomorph level of organization, having no contact between the plates of die flank and those of the ventral shield behind the pectoral fin. It is interchangeable widi Coccosteus spp. and a number of earlier described Gogo forms in a sketch dadogram for arthrodires, but its phylogenedc relationships are unknown. Well preserved material of die scapulocoracoid throws new light on die structure of this element in arthrodires, particularly widi respect to variations in die pattern of canals and foramina.     

A new species of eastmanosteid arthrodire (Pisces: Placodermi) from Gogo, Western Australia

A new species of Eastmanosteus Obruchev is described from the Upper Devonian Gogo Formation of Western Australia as E. calliaspis sp. nov. It is at the pachyosteomorph level of organization having no contact between the plates of the flank and those of the ventral shield behind the pectoral fin. Well preserved material of the parasphenoid and scapulocoracoid throws further light on the structure of these elements in arthrodires. The genus Eastmanosteus is reviewed.     

Study on the systematic position of Metriini based on characters of the larval head (Coleoptera: Carabidae)

Abstract. Characters of the head of larvae of Metrius contractus Eschscholtz, Ozaenini and Paussini are interpreted phylogenetically. The monophyly of Metriini + Ozaenini + Paussini is substantiated by several synapomorphies such as hyperprognathism and strong constriction of the neck. Ozaenini and Paussini together form the sister-group of Metriini. Ozaenini are paraphyletic. The monophyly of Paussini + Ozaenini excluding Pachyteles is indicated by two possible synapomorphies. Several synapomorphies are shared by Physea + Paussini. Secondary prognathism, large membranous submento-mental area and other derived features are considered autapomorphies of Paussini. Paussini excluding Platyrhopalopsis are characterized by the loss of the palpifer. The monophyly of a group which comprises Geadephaga excluding Trachypachini is suggested by several synapomorphic features. A very basal position of the metriine—paussine lineage within Carabidae is indicated by several plesiomorphic features. A hypopharyngeal filter apparatus with a dense fringe of well-arranged, long hairs is a possible autapomorphy of Anisochaeta. The results of this study do not indicate a close relationship between the metriine—paussine lineage and the tribes Brachinini and Crepidogastrini as has been suggested in recent works.     

Beyrichosteus radiatus n. g., n. sp., ein brachythoraker Arthrodire mit vollständig verknöchertem Endokranium aus dem höheren Mittel-Devon (Givetium) der Eifel

A completely perichondrally ossified endocranium of a generalized reprsentative of the Eubrachythoraci,Beyrichosteus radiatus n. g., n. sp., is described. The specimen shows a typical platybasie skull with widely separated nasal capsules like that of the mainly Early Devonian Dolichothoraci. This is contrary to the spezialized generaTapinosteus andPholidosteus from Wildungen, being previously often regarded as model for the endocranium of coccosteomorph arthrodires. However, characters of the orbit and the nasal capsule are more similar to other Eubrachythoraci than to those of Dolichothoraci and primitive Brachythoraci. Until now, there are not enough other examples of sufficiently preserved endocrania of brachythoracid arthrodires known for a phylogenetic analysis. Nevertheless, the observations made onBeyrichosteus are questioning the assignment of the Pholidosteidae to the Coccosteomorphi.     

Monophyly and affinities of albanerpetontid amphibians (Temnospondyli; Lissamphibia)

The Albanerpetontidae are Middle Jurassic-Miocene amphibians that have variously been regarded as caudates (salamanders), a clade distinct from caudates, or incertae sedis lissamphibians. Here I test for monophyly of the Albanerpetontidae and examine the affinities of the group, within the framework of a more inclusive Temnospondyli, by performing a cladistic analysis using 59 informative characters scored for four non-lissamphibian temnospondyl genera, stem- and crown-clade caudates, salientians (frogs), gymnophionans (caecilians), and the two recognized albanerpetontid genera Albanerpeton and Celtedens . Monophyly of the Albanerpetontidae is corroborated. I interpret synapomorphies of the marginal teeth (non-pedicellate; crowns chisel like, labiolingually compressed, with three mesiodistally aligned cuspules) in albanerpetontids as being associated with a shearing bite. Other synapomorphies evidently strengthened and increased the mobility of the skull, mandible, and cervical region for burrowing, feeding, or both. Nested sets of synapomorphies place the Albanerpetontidae within the Lissamphibia, as the sistertaxon of Caudata plus Salientia. None of the 17 characters previously advanced as albanerpetontid-caudate synapomorphies convincingly places the Albanerpetontidae within the Caudata or allies the two groups as sistertaxa. Albanerpetontids are better interpreted not as aberrant caudates, but as a distinct clade of lissamphibians in which numerous apomorphies are superimposed upon an otherwise primitive lissamphibian body plan.     

A cladistic analysis of the genera in the subfamily Pudicinae (Nematoda, Trichostrongyloidea, Heligmonellidae).

A parsimony analysis was performed on 37 specific taxa belonging to the subfamily Pudicinae (family Heligmonellidae), which contains parasites mainly from South American caviomorph rodents. Thirteen characters were used from the synlophe (rotation of axis, presence of carene, carene asymmetry, presence of comaretes, single ventral comarete length, ridge discontinuity, ventral ridge numbers, presence of a peculiar posterior synlophe, presence of supernumerary spines) and the male caudal bursa (relative length of rays 9 and 10, caudal bursa type, division of the dorsal ray, divergence of the 10th rays). The cladogram shows a consistency index of 1.0. The subfamily Pudicinae has two synapomorphies. Two suprageneric groups are recognized. Suprageneric group 1 shows one synapomorphy and contains Heligmostrongylus, Fuellebornema, Sciurodendrium and Pseudoheligmosomum; suprageneric group 2 shows two synapomorphies and contains Pudica, Acanthostrongylus, Justinema and Durettestrongylus. Five genera are defined on the basis of synapomorphies. The genera Heligmostrongylus, Sciurodendrium and Pudica which are considered paraphyletic, however, are retained due to lack of knowledge as to their relationships.     

Metabolism during flight in two species of bats, Phyllostomus hastatus and Pteropus gouldii.

The energetic cost of flight in a wind-tunnel was measured at various combinations of speed and flight angle from two species of bats whose body masses differ by almost an order of magnitude. The highest mean metabolic rate per unit body mass measured from P. hastatus (mean body mass, 0.093 kg) was 130.4 Wkg-1, and that for P. gouldii (mean body mass, 0.78 kg) was 69.6 Wkg-1. These highest metabolic rates, recorded from flying bats, are essentially the same as those predicted for flying birds of the same body masses, but are from 2.5 to 3.0 times greater than the highest metabolic rates of which similar-size exercising terrestrial mammals appear capable. The lowest mean rate of energy utilization per unit body mass P. hastatus required to sustain level flight was 94.2 Wkg-1 and that for P. gouldii was 53.4 Wkg-1. These data from flying bats together with comparable data for flying birds all fall along a straight line when plotted on double logarithmic coordinates as a function of body mass. Such data show that even the lowest metabolic requirements of bats and birds during level flight are about twice the highest metabolic capabilities of similar-size terrestrial mammals. Flying bats share with flying birds the ability to move substantially greater distance per unit energy consumed than walking or running mammals. Calculations show that P. hastatus requires only one-sixth the energy to cover a given distance as does the same-size terrestrial mammal, while P. gouldii requires one-fourth the energy of the same-size terrestrial mammal. An empirically derived equation is presented which enables one to make estimates of the metabolic rates of bats and birds during level flight in nature from body mass data alone. Metabolic data obtained in this study are compared with predictions calculated from an avian flight theory.     

Phylogenetic position of the family Trichodontidae (Teleostei: Perciformes), with a revised classification of the perciform suborder Cottoidei

The phylogenetic relationships of the family Trichodontidae and suborder Cottoidei (and zoarcoid Bathymasteridae) are reconstructed morphologically. The monophyly of the Trichodontidae, Cottoidei, and Zoarcoidei is unambiguously supported by 14 synapomorphies, including 1 newly recognized (and also 4 synapomorphies when ACCTRAN is accepted). It is assumed that the Trichodontidae is deeply nested within the Cottoidei, and the family and Cottoidea plus Cyclopteroidea have a sister relationship, supporting a previously inferred molecular phylogenetic hypothesis. We propose that the Trichodontidae is placed into the cottoid superfamily Trichodontoidea.