Analysis of a Carboniferous embryonic ammonoid assemblage implications for ammonoid embryology

An embryonic ammonoid assemblage was discovered in a carbonate concretion recovered from a dysoxic, relatively offshore marine shale of Virgilian (Upper Pennsylvanian) age in Kansas, USA. The assemblage consists primarily of two species of the Goniatitina, Aristocerassp. and Vidrioceras sp., whose initial chambers (protoconchs) differ in size and shape. Microscopic observations of serial thin sections of specimens at different growth stages reveal the sequence of embryonic shell development starting with the formation of the initial chamber and ending with the synchronous secretion of a prismatic proseptum and nacreous swelliig (primary varix) at the aperture. The mode of occurrence of the embryonic shells of the two species in the concretion suggests that these ammonoids produced numerous small offspring, a reproductive strategy similar to that in many extant coleoids. □ Ammonoids, embryonic shells, development, Carboniferous, Kansas.     

Septal fracture in Nautilus: implications for cephalopod paleobathymetry

Analysis of septal geometry, as embodied in the septal strength index model developed by Westermann (1973, Lethaia 6). , has become a prime avenue for estimating living depths of fossil cephalopods. We have examined the fracturing of Nautilus septa, and its bearing on strength index, by inducing septal rupture under the action of hydrostatic pressure. We found that: (1) septa which actually fail under pressure are not generally weakest as defined by their strength indices; (2) septal strength as defined by the strength index is not correlated with rupture pressure; and (3) most instances of septal failure originate in septal sutures, not in the septa. These results indicate that: (1) septal strength index does not yield wholly reliable strength or depth estimates; and (2) the shortcomings of the strength index model stem from its inability to account for complexities of mechanical failure in morphologically complex cephalopod shells.     

Key innovations in Mesozoic ammonoids: the multicuspidate radula and the calcified aptychus

A nearly complete radula with seven elements per row preserved inside of an isolated, bivalved, calcitic lower jaw (= aptychus) of the Late Jurassic ammonite Aspidoceras is described from the Fossillagerstätte Painten (Bavaria, southern Germany). It is the largest known ammonite radula and the first record for the Perisphinctoidea. The multicuspidate tooth elements (ctenodont type of radula) present short cusps. Owing to significant morphological differences between known aptychophoran ammonoid radulae, their possible function is discussed, partly in comparison with modern cephalopod and gastropod radulae. Analogies between the evolution of the pharyngeal jaws of cichlid fishes and the ammonoid buccal apparatus raise the possibility that the evolution of a multicuspidate radula allowed for a functional decoupling of the aptychophoran ammonoid jaw. The radula, therefore, represents a key innovation which allowed for the evolution of the calcified lower jaws in Jurassic and Cretaceous aptychophoran ammonites. Possible triggers for this morphological change during the early Toarcian are discussed. Finally, we hypothesize potential adaptations of ammonoids to different feeding niches based on radular tooth morphologies.     

Camouflage patterns in Nautilus, and their implications for cephalopod paleobiology

Cowen, R., Gertman, R. & Wiggett, Gail: Camouflage patterns in Nautilus , and their implications for cephalopod paleobiology.
Formal analysis of the pigment patterns of adult Nautilus shows that they are perfectly camouflaged for life in open water. But because of their accretionary growth pattern, juveniles are not fully camouflaged for open water: this supports previous suggestions that young Nautilus are benthonic. The principles of camouflage are used to re-assess some facets of cephalopod paleobiology. The life orientation of some early cyrtocones is re-interpreted. 'Ornament' on ammonoids is seen as camouflage structure, and we infer photic-nektonic, photic-benthonic, and aphotic habitats for three major morphological groups of ammonites. Sexual dimorphism in ammonites was probably accompanied by sexual separation in habitat, except for a short (annual) mating season. This is not inconsistent with modern cephalopod biology.     

Experimental fragmentation patterns of modern Nautilus shells and the implications for fossil cephalopod taphonomy

Shells of modern Nautilus pompilius from the Philippines were experimentally fragmented designed to mimic: (1) transport with sediment; (2) sediment loading; and (3) collision during floating. The breaking patterns by other mechanisms (predation and implosion by hydrostatic pressure) documented in the literature were also considered. The breaking patterns produced by various mechanisms are distinct and can therefore be differentiated. The results allow identification of the distinct mechanisms responsible for specific fragmentation not only in modern Nautilus but also in fossil cephalopods. This is a new approach for the more complete recognition of post-mortem transport of fossil cephalopods and their early taphonomic history, and contributes to our understanding of their palaeobiogeography and palaeoecology.     

The mode of life in ammonoids

The following structural features clearly indicate that ammonoid shells were adapted to withstand considerably higher hydrostatic pressures thanNautilus shells: (1) the corrugated and marginally fluted septa gave the shell wall efficient support against implosion; (2) the secondary connecting rings could grow a great deal in thickness; and (3) the last formed chambers remained full of liquid which supported the last septum. On the basis of the following characters it is concluded that ammonoids were incapable of swimming efficiently by jet-propulsion: (1) the retractor muscles were weakly developed; (2) the life position was unstable and highly variable; and (3) in animals with a ventral apertural rostrum the hyponome was probably absent. Ammonoids are considered here as having been pelagic cephalopods which lived in the upper 1000 m of the oceans, and which probably undertook considerable diurnal vertical migrations, similar to those inSpirula. Only some groups may have adopted a life in shallow epicontinental seas. In the late Mesozoic, ammonoids have been replaced by modern oceanic squids which are extremely numerous in the corresponding pelagic environment.     

Tooth function and replacement in early Mesozoic ornithischian dinosaurs: implications for aestivation

Thulborn (1978, Leihaia II ) suggests that ornithischian dinosaurs of the upper Stormberg Series (Late Triassic-Early Jurassic) of southern Africa underwent aestivation during an annual dry season. His argument, based on an interpretation of tooth function and replacement in heterodontosaurids, is: (1) unequivocal evidence of tooth replacement is not seen, and (2) piecemeal replacement of the dentition would be incompatible with maintenance of a fore-aft grinding function of the teeth; therefore, the entire dentition must have been rapidly replaced as a unit during periods of non-feeding, i.e. during aestivation. However, study of tooth wear patterns in Lanasaurus, Lycorhinus , and Heterodontosaurus show that jaw movements during mastication were orthal (open-and-close) and lateral to medial, not forwards and backwards. Differences in degree of tooth wear would not interfere with masticatory movements. Patterns of differential wear indicate that tooth replacement was not periodic but continuous, as in other reptiles. Zahnreihen , with a Z-spacing of about 3.0, are recognizable. Replacement ceased in mature individuals. The dentition shows adaptations for prolonging its effective life despite heavy wear. Differential tooth wear is incompatible with the idea of replacement of the entire dentition as a unit during an hypothesized period of aestivation. Thulborn's suggestion of aestivation in fabrosaurid ornithischians is also shown to be unlikely.     

Mesozoic woods from India: Nomenclature review and palaeoclimatic implications

Nomenclature reappraisal, diversity pattern and palaeoclimatic implications of Jurassic, Triassic and Early Cretaceous pycnoxylic woods in India are undertaken in the present study. Among the fourteen generic names published previously, only eight are validly published and the rest are nomenclaturally illegitimate. About 51 species were reported under these genera to date. There is a gradual increase of species diversity of fossil wood from the Triassic to Early Cretaceous. The nature of the growth rings was applied to understand the palaeoclimate. The lack of distinct growth rings in the Triassic woods suggests absence of seasonality. The Jurassic woods with an inconsistency in growth rings and presence of growth interruptions suggest climate was seasonal and turbulent. During the Early Cretaceous, conifer dominated vegetation and with wider growth rings and gradual transition suggests warm environments with pronounced seasonality. The general increase in mean ring width from the Triassic to Early Cretaceous indicates ameliorating climatic conditions, particularly benign summer conditions.     

Nautilus at risk--estimating population size and demography of Nautilus pompilius

The low fecundity, late maturity, long gestation and long life span of Nautilus suggest that this species is vulnerable to over-exploitation. Demand from the ornamental shell trade has contributed to their rapid decline in localized populations. More data from wild populations are needed to design management plans which ensure Nautilus persistence. We used a variety of techniques including capture-mark-recapture, baited remote underwater video systems, ultrasonic telemetry and remotely operated vehicles to estimate population size, growth rates, distribution and demographic characteristics of an unexploited Nautilus pompilius population at Osprey Reef (Coral Sea, Australia). We estimated a small and dispersed population of between 844 and 4467 individuals (14.6-77.4 km(-2)) dominated by males (83:17 male:female) and comprised of few juveniles (<10%).These results provide the first Nautilid population and density estimates which are essential elements for long-term management of populations via sustainable catch models. Results from baited remote underwater video systems provide confidence for their more widespread use to assess efficiently the size and density of exploited and unexploited Nautilus populations worldwide.     

Size scaling and stiffness of avian primary feathers: implications for the flight of Mesozoic birds

The primary feathers of birds are subject to cyclical forces in flight causing their shafts (rachises) to bend. The amount the feathers deflect during flight is dependent upon the flexural stiffness of the rachises. By quantifying scaling relationships between body mass and feather linear dimensions in a large data set of living birds, we show that both feather length and feather diameter scale much closer to predictions for geometric similarity than they do to elastic similarity. Scaling allometry also indicates that the primary feathers of larger birds are relatively shorter and their rachises relatively narrower, compared to those of smaller birds. Two-point bending tests indicated that larger birds have more flexible feathers than smaller species. Discriminant functional analyses (DFA) showed that body mass, primary feather length and rachis diameter can be used to differentiate between different magnitudes of feather bending stiffness, with primary feather length explaining 63% of variance in rachis stiffness. Adding fossil measurement data to our DFA showed that Archaeopteryx and Confuciusornis do not overlap with extant birds. This strongly suggests that the bending stiffness of their primary feathers was different to extant birds and provides further evidence for distinctive flight styles and likely limited flight ability in Archaeopteryx and Confuciusornis.     

Carbon isotopic abundances in Mesozoic and Cenozoic fossil plants: Palaeoecological implications

Carbon isotopic abundances have been measured for more than one hundred samples of fossil plants ranging in age from middle Triassic to late Tertiary. Most of the plant fossils were identified at the specific or generic level and were selected as representing a variety of continental environments, including xeric and humid habitats. Material analysed included numerous fragments of flowers, seeds, fruits, leaves and wood, as well as a single amorphous lignite sample. The analyses performed for the plant fragments indicate relatively constant isotopic compositions during this time interval, with plant δ¹³C values ranging between -28 and -20%. These values are within the range for living terrestrial plants with C₃, photosynthesis, although values more positive than -23% are rare in C₃ plants and typically found in plants growing under environmental stress. Lower δ¹³C values might have been expected owing to the much higher CO₂, levels of the Cretaceous atmosphere that have been inferred from marine carbonates. No fossils with values indicating C₄, photosynthesis were discovered. Fossil plants from inferred mesic environments showed δ¹³C values ranging between -26.7 and -24.1%. Highest δ¹³C values in angiosperms (up to -20.1%) were measured for Late Cretaceous combretaceous flowers from Portugal. Some cheirolepidiaceous conifers from the Early Cretaceous also showed high δ¹³C values. Values measured for Pseudofrenelopsis varians and Glenrosa taxensis were -21.9%, and values of gymnosperm wood, probably of cheirolepidiaceous affinity, were -19.0%. These high values are in accordance with inferred ecological conditions for the fossil plants. They may suggest a tendency for C₄,-like photosynthesis, although the data are equivocal. Higher values (-17.3%) clearly falling outside the C₃, range were, however, obtained from a single lignite fragment of Late Cretaceous (Maastrichtian) age. The nature of this plant fragment is unknown, but the result suggests that C₄-like photosynthesis was present at least in some latest Cretaceous vegetation. A hadrosaurian dinosaur with well-preserved collagen-like organic matter from the same deposit showed δ¹³C values around-16%, which also suggests the presence of CAM or even C₄ plants in the latest Cretaceous. □Carbon isotopic abundances, δ¹³C values, dinosaurs, plants, photosynthetic pathways, Mesozoic.     

The biogeography of Early Triassic ammonoid faunas: Clusters, gradients, and networks

The aim of this paper is to quantitatively investigate the spatial and temporal biogeographical relationships of the recovery of ammonoid faunas after the Permian-Triassic mass extinction using three complementary numerical approaches among which is a new, non-hierarchical clustering strategy. The faunal data set consists of a taxonomically homogenised compilation of the spatial and temporal occurrences of ammonoid genera within 20 Early Triassic Tethyan and Panthalassic sites ranging from 40°S to 70°N in palaeolatitudes. In addition to hierarchical cluster analysis (hCA) and nonmetric multidimensional scaling (NMDS), we introduce a third, new non-hierarchical clustering technique allowing the visualisation of a nonmetric interassemblages similarity structure as a connected network constructed without inferring additional internal nodes. The resulting network, which we call a “Bootstrapped Spanning Network” (BSN), allows the simultaneous identification of partially or totally nested as well as gradational linear or reticulated biogeographical structures.The identified interlocalities relationships indicate that the very beginning of the Early Triassic (Griesbachian) corresponds to a very simple biogeographical context, representing a time of great cosmopolitanism for ammonoids. This context shifts rapidly to a more complex configuration indicative of a more endemic and latitudinally-restricted distribution of the ammonoids during the middle and late Early Triassic (Smithian and Spathian). From an evolutionary dynamic point of view, our results illustrate a very rapid (less than ca. 1.4 myr) Early Triassic recovery of the ammonoid faunas, in contrast to many other marine organisms. This recovery is linked with a marked increase in the overall biogeographical heterogeneity, and parallels the formation of a latitudinal gradient of taxonomic richness, which may be essentially controlled by the progressive intensification of the gradient of sea surface temperature. From a methodological point of view, we show that a BSN is a simple, intuitively legible picture of the nested as well as gradational taxonomic similarity relationships, hence providing a good synthesis (and additional insights) between hierarchical clustering and ordination in reduced space results.     

Oligochitosan derivatives bearing electron-deficient aromatic rings for adsorption of amitriptyline: implications for drug detoxification

The objective of this work is the synthesis of water-soluble oligochitosan derivatives with electron deficient aromatic rings for selective and rapid adsorption of amitriptyline through pi-pi complexation. Oligochitosan was chemically modified under homogeneous conditions in dimethyl sulfoxide (DMSO). (1)H NMR, FT-IR, and MALDI-TOF were employed in characterization, confirming that the electron deficient aromatic rings were chemically attached to the backbone of oligochitosan. Thromboelastography (TEG) revealed functionalized oligochitosan derivatives did not affect blood clotting. (1)H NMR was also utilized to observe the aromatic-aromatic interaction between electron deficient aromatic rings on oligochitosan and electron rich aromatic rings in amitriptyline. The chemical shift variation of aromatic protons in oligochitosan derivatives was followed to monitor the aromatic-aromatic interaction. Upfield shift of aromatic protons on benzenesulfonyl and dinitrobenzenesulfonyl groups was observed upon the addition of amitriptyline, supporting the formation of pi-pi complexes through aromatic-aromatic interactions. Dinitrobenzenesulfonyl rings show a larger variation in chemical shift due to the presence of the electron deficient nitro groups.     

Multivariate analysis of neognath skeletal measurements: implications for body mass estimation in Mesozoic birds

The abundant fossils of avian stem taxa unearthed during the last years make it necessary to review and improve the models for estimating body mass used in palaeoecological studies. In this article, single and multiple regression functions based on osteological measurements were obtained from a large data set of extant flying birds for estimating the body mass of 42 Mesozoic specimens from stem taxa Archaeopterygidae, Jeholornithidae, Sapeornithidae, Confuciusornithidae, and Enantiornithes, and basal members of Ornithuromorpha. Traditionally, body mass has been estimated in fossil vertebrates using univariate scaling functions. In contrast, multiple regression functions have been used less frequently. Both predictive methods can be affected by different sources of error from statistics, phylogenetic relationships, ecological adaptations, and bone preservation; however, although some studies have addressed these biases, few have tested them within the context of a single data set. In our data set, we find that the models with greater predictive strength and applicability for new specimens, especially for stem taxa, are those derived from multiple regression analyses. For this reason, we suggest that multiple regression analyses may provide improved predictive strength for stem group specimens. Moreover, the methodology used for selecting variables allowed us to obtain specific sets of predictors for each fossil stem group that presumably minimized the variation resulting from historical contingency (i.e. differences in skeletal morphology arising from phylogeny), locomotor adaptations, and diagenetic compaction. The loss of generalizability in the multiple regression models resulting from collinearity effects was negligible on the body mass estimates derived from our data set. Therefore, the body mass values obtained for Mesozoic specimens are accurate and can be used in future studies in a number of palaeobiological and evolutionary aspects of extinct birds, particularly the first stages of avian flight. © 2015 The Linnean Society of London     

The genus Luppovia and the phylogeny of Cretaceous heteromorphic ammonoids

The unique preservation of heteromorphic shells of Luppovia Kakabadze, Bogdanova & Mikhailova 1978 from the Aptian deposits of Bolshoj Balkhan (western Turkmenia) made it possible to study the microstructure, internal shell characteristics and suture ontogeny of this genus. Microstructural investigations revealed that the dorsal shell wall in Luppovia is composed of three layers in contrast to that of monomorphic ammonoids, which is single-layered. This ability of secreting different numbers of layers in monomorphs and heteromorphs demonstrates plasticity of secreting activity of the mantle epithelium. Data obtained from the study of internal shell structure and suture ontogeny have been compared. Both these methods of investigation lead to the conclusion that the genus Luppovia belongs to the order Ammonitida. The results obtained confirm the viewpoint that the Cretaceous heteromorphic superfamilies Turrilitaceae and Ancylocerataceae have independent origins and belong to two different orders, Lytoceratida and Ammonitida, respectively. □ Ammonoidea , Luppovia, shell structure, ontogeny, phylogeny, Cretaceous.     

Morphology and histology of the digestive tract of Nautilus pompilius and Nautilus macromphalus (Cephalopoda, Tetrabranchiata)

 This study presents histological and scanning electron microscopical findings on the structural differentiation, and the nervous and vascular supply of the digestive tracts of Nautilus pompilius and N. macromphalus, including the foregut, stomach, vestibulum, caecum, midgut and rectum. The stereoscopic reconstruction of the vestibulocaecal complex gives an idea how the digestive cycle between the stomach, vestibulum, caecum and proximal midgut could possibly proceed. All parts of the digestive tract are covered luminally by a columnar epithelium which contains numerous goblet cells. The epithelium is ciliated in the vestibulum, caecum, proximal midgut and the longitudinal groove of the rectum. On this lamina epithelialis mucosae borders the lamina propria mucosae, which consists of connective tissue and some muscle cells. In the stomach it is differentiated, forming a special bolster-like layer. The lamina propria mucosae is followed by the tunica muscularis, which consists of a stratum circulare and a stratum longitudinale in the foregut, vestibulum, caecum, midgut and rectum. In the stomach, midgut and rectum, the tunica adventitia, which consists of a thin layer of connective tissue, is located between the tunica muscularis and the cuboidal tunica serosa. Accepted: 4 August 1997