AMMONOID SHELL STRUCTURES OF PRIMARY ORGANIC COMPOSITION

Abstract:  Palaeozoic and Mesozoic cephalopod conchs occasionally reveal dark organic coatings at the aperture. A number of these coatings, including still unrecorded examples, are described, figured and interpreted herein. On the basis of elemental analysis, actualistic comparison and a comparison with Triassic bivalves, some of these coatings are shown to consist of apatite and primarily probably of conchiolin (and also probably melanin). In several Mesozoic ammonoid genera such as Paranannites , Psiloceras , Lytoceras , Phylloceras , Harpoceras and Chondroceras , some of these coatings (recorded herein for most of these taxa for the first time) are interpreted as a structure similar to the black band, which was previously known only from Recent Allonautilus and Nautilus . In contrast to these nautilid genera, however, the organic material of some Mesozoic ammonoids was not deposited on the inside of the shell but externally, albeit positioned at the terminal aperture as in Recent nautilids. Some ammonoids of Carboniferous and Triassic age show several such bands at more or less regular angular distances on the ultimate whorls and at the aperture, e.g. Nomismoceras , Gatherites , Owenites , Paranannites , Juvenites and Melagathiceratidae gen. et sp. nov. Triassic material from Oman shows that the black coating was probably secreted from the inside, because the position of this organic deposit changes from interior to exterior in an anterior direction (i.e. adaperturally). This structure has previously been referred to as a 'false colour pattern' and is here interpreted as having been formed at an interim aperture or megastria ('alter Mundrand'). All structures discussed in the paper are considered to have been secreted by a single organ and to have been initiated by some form of stress or adverse conditions. Thus, certain environmental parameters and growth anomalies appear to have influenced their formation.     

Flexible nacre in the nautiloid Isorthoceras, with remarks on the evolution of cephalopod nacre

Secondarily phosphatized shells of Isorthoceras sociale (Hall) from the Upper Ordovician Maquokcta shale, Iowa, U.S.A., often have their original ultrastructurc preserved. Scanning Electron Microscope study reveals that the nacreous layer is similar to that in the shells of Recent Nautilus , except that the nacreous tablets usually have a much larger diameter, and that numerous vertical interspaces arc left between adjacent stacks of tablets. Because of the latter condition, the nacre in A sociale is porous, whereas it is compact in Nautilus. The pores in the nacre were probably filled with a non-mincralizcd organic matrix. The high porosity and high content of organic matrix made the nacreous layer of I. sociale much more flexible and considerably less strong than that in Nautilus. As a consequence, the shells of I. sociale , and probably also of other orthoconic cephalopods, were more flexible and could not withstand such high hydrostatic pressures as the shells of Nautilus. During evolution, the porosity of the nacreous layer became completely reduced. As a result, the mechanical properties of the nacre changed towards a lower flexibility and a higher tensile strength. Cephalopoda, Nautiloidea, nacre, ultrastructurc, phosphatization, Upper Ordovician.     

Occluded Umbilicus In The Pinacitinae (Devonian) And Its Palaeoecological Implications

Eifelian (Middle Devonian) ammonoids of the Pinacitinae Hyatt, 1900 ( Exopinacites , Pinacites )with preserved shell structures from the eastern Anti–Atlas (Morocco) have revealed unusual morphological features. The Pinacitinae belong to the earliest ammonoids which closed their umbilici. As an approach to an interpretation of these structures, the representatives of the subfamily Pinacitinae ( Exopinacites singularis , Pinacites jugleri , P. eminens ) are compared with other ammonoids, e.g. Acrimeroceras , Araucanites , Clistoceras , Gaudryceras , Nathorstites , Prolobites , and Synpharciceras , which produced umbilical plugs and covers. Some of these are comparable in structure to Nautilus pompilius and N. belauensis . In contrast to all of these taxa, the lateral shell wall of the Pinacitinae reached the centre of the umbilicus and formed an umbilical lid. The umbilical shell wall rests on the umbilical lid of the previous whorl. This construction probably had the advantage that it improved the hydrodynamic properties of the conch, along with the oxyconic conch shape and the approximately horizontal orientation of the aperture.     

Soft‐tissue imprints in fossil and Recent cephalopod septa and septum formation

Several soft‐tissue imprints and attachment sites have been discovered on the inside of the shell wall and on the apertural side of the septum of various fossil and Recent ectocochleate cephalopods. In addition to the scars of the cephalic retractors, steinkerns of the body chambers of bactritoids and some ammonoids from the Moroccan and the German Emsian (Early Devonian) display various kinds of striations; some of these striations are restricted to the mural part of the septum, some start at the suture and terminate at the anterior limit of the annular elevation. Several of these features were also discovered in specimens of Mesozoic and Recent nautilids. These structures are here interpreted as imprints of muscle fibre bundles of the posterior and especially the septal mantle, blood vessels as well as the septal furrow. Most of these structures were not found in ammonoids younger than Middle Devonian. We suggest that newly formed, not yet mineralized (or only slightly), septa were more tightly stayed between the more numerous lobes and saddles in more strongly folded septa of more derived ammonoids and that the higher tension in these septa did not permit soft‐parts to leave imprints on the organic preseptum. It is conceivable that this permitted more derived ammonoids to replace the chamber liquid faster by gas and consequently, new chambers could be used earlier than in other ectocochleate cephalopods, perhaps this process began even prior to mineralization. This would have allowed faster growth rates in derived ammonoids.     

On the efficiency of the buoyancy apparatus in ammonoids: evidences from sublethal shell injuries

The five greatest sublethal injuries were selected from a collection of more than 12,000 predominantly Mesozoic injured or otherwise pathological ammonoids. The loss of shell mass from these survived injuries was calculated and compared with comparable tolerances in the recent Nautilus . These ammonoids tolerated a shell loss up to four times greater than in Nautilus . The maximum tolerated shell loss indicates an unexpected buoyancy compensation mechanism. The buoyancy of the selected specimens was calculated. The results show that the buoyancy of all the observed ammonoid shells was positive. In order to maintain neutral buoyancy after injury, these ammonoids had to fill the phragmocone with a volume of mass. Nautilus compensated a maximum mass loss requiring a liquid refill of 3% of the cameral capacity, the ammonoids compensated a maximum of observed mass loss requiring a liquid refill of more than 10% of cameral capacity. The ratio of chamber volume/siphuncular surface area in the ammonoid Lithacoceras is 0.043, indicating that the relative area of the siphuncular epithelium in Lithacoceras is significantly higher when compared with a ratio of 0.12-0.14 in the adult Nautilus . The phragmocone in ammonoids offered the ability of a much more active buoyancy regulation than in Nautilus .     

Nautilus—a poor model for the function and behavior of ammonoids?

Inferences drawn from the biology, function, and behavior of closely related living forms facilitate interpretation of the mode of life of groups known only from the fossil record. The choice of phylogenetically relevant modern 'model organisms' can have critical bearing on the resulting interpretations. The biology and behavior of fossil ammonoids are often interpreted in the light of evidence derived from the study of modern Nautilus . However, examination of the fossil record and cladistic analyses both indicate that coleoids are much more closely related to ammonoids than is Nautilus . Coleoid biology and behavior differ dramatically from the biology and behavior of Nautilus . Thus, the inclusion of coleoids as examples, rather than reliance on Nautilus alone, produces a strikingly different vision of ammonoid biology and suggests that inferences of ammonoid biology and behavior that rely exclusively on Nautilus should be reviewed. Two features related to swimming ability in Nautilus , static stability and large retractor muscles, are much reduced in many ammonoids, leading to the interpretation that ammonoids were poorer swimmers than Nautilus . However, reexamination of the evidence indicates that static stability should not play a role in the swimming of ammonoids with long body chambers. In addition, functional arguments suggest that a coleoid-like swimming mechanism should have evolved prior to the loss of the body chamber in coleoids. Thus, a coleoid-like swimming mechanism is likely to have evolved prior to the separation of ammonoid and coleoid lineages. A mechanism is proposed by which a coleoid swimming mechanism, independent of retractor muscle size, could function in ammonoids with long body chambers.□ Ammonoids, ammonites, evolution, functional morphology , Nautilus, phylogeny .     

Biological response to experimental damage of the phragmocone and siphuncle in Nautilus pompilius Linnaeus

Tsujino, Y & Shigeta, Y. 2012: Biological response to experimental damage of the phragmocone and siphuncle in Nautilus pompilius Linnaeus. Lethaia, Vol. 45, pp. 443–449. Three adult specimens of Nautilus pomplilius Linnaeus from the Philippines were experimented on to estimate the biological response to damage of the phragmocone and siphuncle in this cephalopod mollusc. In addition, the data obtained from the experiments were used for discussion of shell damage in ammonoids and in other extinct cephalopods. Specimen’s phragmocone and siphuncle were perforated and severed artificially, followed by observations in the laboratory tank during periods of 75 and 132 days. For at least 2 or 3 months, all individuals survived after damage to the phragmocone and siphuncle despite loss of neutral buoyancy. Based on our observations after completion of the experiments, the severed adoral remaining part of siphuncle healed by the siphunclar epithelium. In addition, perforation of the phragmocone was partly repaired by shell secretion from the dorsally extending mantle due to subsequent volution of shell growth. Our experiments revealed that damage to the phragmocone and siphuncle in Nautilus was not necessarily a lethal injury. It may be possible that such biological response also applies to extinct ammonoids and nautiloids. In a similar case of extinct ammonoids and nautiloids, damage to their phragmocone and siphuncle may also not have been a lethal injury as with Nautilus. However, some factors leading to death are likely to be dependent on the degree of damage to the phragmocone and siphuncle and influence of hydraulic pressure. □Ammonoids, injury, nautiloids, Nautilus, phragmocone, repair, siphuncle.     

Rhyncholites and conchorhynchs as calcified jaw elements in some late Cretaceous ammonites

Conspicuous calcareous coverings are present in the anterior region of 17 fossil jaws from late Cretaceous rocks of Hokkaido (Japan) and Sakhalin (U.S.S.R.). The jaws were preserved in calcareous nodules either in situ in body chambers of ammonites or in close association with identifiable ammonite conch remains. From the morphologic similarity between in situ and isolated jaws, they may be attributed to Tetragonites glabrus, Gaudryceras tenuiliratum, G. denseplicatum, G. sp., and Neophylloceras subramosum. The jaw apparatus of these species is composed of two three-dimensional black walls of carbonate apatite, which might be a diagenetic replacement of chitinous material. The calcareous coverings in both upper and lower jaws closely resemble those of upper (rhyncholite) and lower (conchorhynch) jaws of modern Nautilus as well as rhyncholite and conchorhynch fossils in their gross morphology, microstructure, and chemical composition. Calcified remains of cephalopod jaws known as rhyncholites and conchorhynchs have been reported from late Paleozoic to Recent. The present discovery of ammonoid rhyncholites and conchorhynchs suggests that at least some previously known late Paleozoic and Mesozoic counterparts belong to the Ammonoidea. The essential similarity of jaw elements of some Late Cretaceous ammonites and modern Nautilus gives reliable information on the feeding habits of the former. The sharp and thick ammonoid rhyncholites and conchorhynchs may have had a special function for cutting up food, similar to those of Nautilus.     

Phylogeny of the aptychi-possessing Neoammonoidea (Aptychophora nov., Cephalopoda)

The different forms of the aptychi (opercula, homologous with lower jaws) of the Ammonoidea are used for the first time in a phylogenetic analysis of part of the classic Ammonoidea phylogeny. The results indicate that the aptychi-possessing ammonoids form a monophylum for which we propose the informal name Aptychophora nov. Among the Jurassic ammonoids, it is possible to recognize several monophyletic groups. In part, our results support existing superfamilies (e.g. Hildocerataceae, Haplocerataceae) by new synapomorphies. However, the Perisphinctaceae can now be much more clearly differentiated than in the previously established phylogenetic tree. The Upper Cretaceous ammonoid superfamilies cannot be derived from the Haplocerataceae, but are descendants of a 'primitive' perisphinctacean possessing a praestriaptychus. Nor can they be derived from the 'higher' perisphinctaceans (family Perisphinctidae) because that clade is characterized by granulaptychi. The consequence of these results is that the quadrilobate primary suture of the 'Ancyloceratina' must have evolved more than once by reduction from an ancestral quinquelobate primary suture. The Ancyloceratidae have praestraptychi or aptychi types which can be derived from praestriaptychi, whereas the Crioceratitinae have longitudinally striated anaptychi.     

Adhesive mechanisms in cephalopods: a review

Several genera of cephalopods (Nautilus, Sepia, Euprymna and Idiosepius) produce adhesive secretions, which are used for attachment to the substratum, for mating and to capture prey. These adhesive structures are located in different parts of the body, viz. in the digital tentacles (Nautilus), in the ventral surface of the mantle and fourth arm pair (Sepia), in the dorsal epidermis (Euprymna), or in the dorsal mantle side and partly on the fins (Idiosepius). Adhesion in Sepia is induced by suction of dermal structures on the mantle, while for Nautilus, Euprymna and Idiosepius adhesion is probably achieved by chemical substances. Histochemical studies indicate that in Nautilus and Idiosepius secretory cells that appear to be involved in adhesion stain for carbohydrates and protein, whilst in Euprymna only carbohydrates are detectable. De-adhesion is either achieved by muscle contraction of the tentacles and mantle (Nautilus and Sepia) or by secretion of substances (Euprymna). The de-adhesive mechanism used by Idiosepius remains unknown.     

Functional interpretation of inner shell layers in Triassic ceratid ammonites

The wrinkle layer the inner prismatic layer are described in three Triassic ceratid genera: Phyllocludiscites. Megaphyllites Proarcestes. Both layers have their counterparts in the shell wall of the recent Nautilus: the wrinkle layer corresponds to the mantle-adhesive layer the inner prismatic layer to the myostracal layer in Nautilus. A detailed structural functional comparison between these layers is given. The wrinkle layer is also compared with the oblique prismatic layer in recent gastropods.     

Phylloceratina ammonoids in the Iberian Basin during the Middle Jurassic: a model of biogeographical and taphonomic dispersal related to relative sea-level changes

Phylloceratina and Lytoceratina ammonoids in the Middle Jurassic of the Iberian Chain (E. Spain) represent less than 1% of the whole of Ammonoidea. There are two intervals, however, in which their occurrence is remarkably constant: within the Upper Bajocian and at the Lower/Middle Callovian boundary. These two dispersal episodes of Phylloceratina and Lytoceratina into the Iberian Basin are regarded to reflect changes in their palaeoecological and taphonomical behaviour, as a consequence of regional sea-level changes during the Middle Jurassic. A relative rise during the Late Bajocian favoured the immigration of juvenile phyllocerataceans. Phylloceras and Adabofoloceras immigrations gave rise to monospecific assemblages, where they soon died. They did not breed or ontogenically develop in this basin. In contrast, phyllocerataceans recorded at the Lower/Middle Callovian boundary constitute polyspecific assemblages, dominated by adult individuals. These Callovian assemblages were formed by nekroplanktonic drift, related to a relative sea-level fall and general homogenization of the shelf bottom, hence favouring the concentration of remains of organisms from more open marine and oceanic areas.     

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.     

Post-hatching early life history of Cretaceous Ammonoidea

Post-hatching early life histories in Cretaceous Ammonoidea are discussed on the basis of density calculations of the shells in 71 species belonging to four separate suborders. The calculation was made under the assumption that a newly hatched ammonoid had a gas-filled chamber and a succeeding body-filled whorl terminating at the primary constriction. The results show that the density of the species examined at the hatching stage is almost constant and is relatively smaller than that of seawater, i.e. the animals are positively buoyant. This fact strongly suggests a planktic mode of life. In all species, the density increases gradually with growth and attains neutral buoyancy at 2.C2.5 mm in shell diameter. Thus, most ammonoids probably changed their mode of life from planktic to nektoplanktic or nektobenthic at this critical point. The rare occurrence of newly hatched specimens (ammonitellas) in many ammonoid assemblages may also support this interpretation. Planktic duration of a newly hatched ammonoid might be regulated by the animal's density at hatching, shell growth pattern, cameral volume (or hatching size), and rate of cameralliquid removal (or siphuncle diameter). The latter two seem to be very important factors in determining the biogeographical framework of species, as demonstrated in the Tetragonitaceae.□ Cretaceous, Ammonoidea, density calculation, early life history .     

The Luoping biota: exceptional preservation, and new evidence on the Triassic recovery from end-Permian mass extinction

The timing and nature of biotic recovery from the devastating end-Permian mass extinction (252 Ma) are much debated. New studies in South China suggest that complex marine ecosystems did not become re-established until the middle–late Anisian (Middle Triassic), much later than had been proposed by some. The recently discovered exceptionally preserved Luoping biota from the Anisian Stage of the Middle Triassic, Yunnan Province and southwest China shows this final stage of community assembly on the continental shelf. The fossil assemblage is a mixture of marine animals, including abundant lightly sclerotized arthropods, associated with fishes, marine reptiles, bivalves, gastropods, belemnoids, ammonoids, echinoderms, brachiopods, conodonts and foraminifers, as well as plants and rare arthropods from nearby land. In some ways, the Luoping biota rebuilt the framework of the pre-extinction latest Permian marine ecosystem, but it differed too in profound ways. New trophic levels were introduced, most notably among top predators in the form of the diverse marine reptiles that had no evident analogues in the Late Permian. The Luoping biota is one of the most diverse Triassic marine fossil Lagerstätten in the world, providing a new and early window on recovery and radiation of Triassic marine ecosystems some 10 Myr after the end-Permian mass extinction.     

Times of crisis in the evolution of the cephalopoda

The evolutionary history of the ectocochlian cephalopods is punctuated by a number of severe crises during each of which this class came very close to extinction. The crisis events follow each other at intervals of from seven to almost 300 million years and, with one exception, were not synchronous for Nautiloidea and Ammonoidea. Only at the end of the Triassic period did both groups simultaneously face the danger of extinction. Generally, the survivors of crisis situations have simple shell forms and are strikingly similar to each other. To trace the details of cephalopod evolution, the family on the taxonomic level and the stratigraphic stage on the chronological level do not provide scales fine enough to reconstruct the true course of this process. The causes of crises and “mass extinctions” are not yet understood. Most authors have approached this problem in a simplistic manner, searching for a single cause for any, or all, events of this kind. It seems that we do not even have begun to understand what the problems are.     

The coarse wrinkle layer of Palaeozoic ammonoids: new evidence from the Early Carboniferous of Morocco

The wrinkle layer is a dorsal shell structure occurring in a number of ammonoids, but its function is still debated. Here, we describe, from Moroccan material of the Early Carboniferous species Maxigoniatites saourensis (Pareyn, 1961 ), the most conspicuous wrinkle layer known within the Ammonoidea. This additional shell layer occurs in the ventrolateral portion of the adult body chamber and forms continuous lamellae, which range about two millimetres into the lumen of the body chamber. Possible functions are discussed and the most likely interpretation for the structure is ‘fabricational noise’, which is related to the coarsening of the shell ornament of the terminal body chamber.     

GUODUNITES, A LOW-PALAEOLATITUDE AND TRANS-PANTHALASSIC SMITHIAN (EARLY TRIASSIC) AMMONOID GENUS

Abstract:  Based on new, bed-rock controlled material from Oman and Utah, USA, the Early Triassic genus Guodunites , which was recently erected on the basis of scarce specimens from northwestern Guangxi, South China, is now shown to be a representative of Proptychitidae. This solves the question of the previously unknown phylogenetic affinity of this genus. The genus is restricted to the late middle Smithian, and to date, its biogeographical distribution comprises Oman, South China and Utah, thus indicating an essentially low palaeolatitudinal distribution during the Early Triassic. Its palaeobiogeographical distribution further strengthens the existence of significant equatorial faunal exchanges between both sides of the Panthalassa at that time. It also suggests that, in addition to the potential stepping stones represented by Panthalassic terranes, vigorous equatorial oceanic currents must have contributed largely to the dispersal of ammonoids during such time intervals.