How to recognize in situ fossil cephalopods: evidence from experiments with modern Nautilus

Field and flume experiments with modern Nautilus pompilius establish two prerequisites to recognize in situ preservation of fossil cephalopod shells (soft parts were within body chamber in situ at the time of fossilization): occurrence of the upper jaw within the body chamber and the position of jaws within the body chamber. Morphology of shells and jaws in modern and fossil nautiloids is so similar that these prerequisites can be applied for fossil nautiloids and provide implications for ammonoids. The upper jaws of Nautilus start to move at a water velocity of > 0.2 m/s, when the shells are reoriented with the aperture downstream; jaws are therefore unlikely to be secondarily deposited near the shell aperture by bottom currents. The lower jaws, moved at the velocity of > 0.1 m/s, can be deposited around the shell aperture by weak current (0.1–0.2 m/s in velocity), but never enter the inside of body chamber. Neither jaw is likely to be separately and selectively displaced from the inside of the body chamber through scavenging of the soft parts by burrowing infaunal animals. An upper jaw preserved inside the body chamber, together with a lower jaw, is thus a reliable indicator of in situ preservation; a sole lower jaw preserved around the shell aperture is likely to be secondarily deposited. Sedimentary structures inferring rapid burial events and jaw size are useful as additional evidence. Smaller jaws were more likely to be displaced from the body chamber by scavenging by infaunal animals after in situ burial, so that smaller jaws preserved within the body chamber suggest less scavenging. These findings are crucial to interpreting the taphonomic history and palaeo-ecology of fossil cephalopods.     

Evidence for Palaeozoic orthoconic cephalopods with bimineralic shells

Based on the aragonite composition of extant and exceptionally preserved fossil cephalopods going back to the early Palaeozoic, it is commonly assumed that all externally shelled cephalopods had an aragonitic shell wall. We demonstrate herein that at least two taxa of Siluro‐Devonian orthoconic nautiloids (Dawsonoceras, Spyroceras) had an original bimineralic shell, which developed convergently with gastropods and bivalves.     

Ammoniten mit Kieferapparat und Radula aus Lias-Geschieben

Jaws and radula have been found in two adult microconch specimens of the Liassic ammonite speciesEleganticeras elegantulum (Young & Bird, 1828) from glacial drift boulders near Hamburg. They are more similar to those of recent dibranchiates than ofNautilus, the radula having seven longitudinal rows of teeth instead of thirteen in recentNautilus. Since bothOwens classification of the cephalopods into Dibranchiata and Tetrabranchiata according to the number of gills (which cannot be observed in fossil cephalopods) andSchwarz’ division into Ecto- and Endocochlia according to the (assumed) relative position of body and shell are rather unfortunate, the radula is suggested as a sound basis of classification. It is acceptable to paleo- as well as to neozoologists. As yet, the known evidence is sufficient for a twofold division only: Class Cephalopoda Subclass Lateradulata: Nautiloidea Subclass Angusteradulata: Ammonoidea Coleoidea     

Shell microstructure in the Permian nonmarine bivalve Palaeomutela Amalitzky: Revision of the generic diagnosis

The microstructure of aragonitic and calcitic shells of the genus Palaeomutela Amalitzky, 1891 is examined. The aragonitic shell consists of three main layers, each is distinguished by certain crossed lamellar microstructure: comarginal, radial, and complex. As aragonite is recrystallized into pelitic calcite, microstructural shell features are preserved. Many species of Palaeomutela from localities of different age display the same microstructural pattern, which is possible to regard as a character of generic rank.     

Double function of aptychi (Ammonoidea) as jaw elements and opercula

Lehmann, U. & Kulicki, C. 1990 10 15: Double function of aptychi (Ammonoidea) as jaw elements and opercula. Lethaia , Vol. 23, pp. 325–331. Oslo. ISSN 0024–1164.
Aptychi are calcitic coverings on the outer surface of organic ammonite lower jaws. They are similar in shape to that of the corresponding ammonite apertures. This observation and additional features of many aptychi are in harmony with their former interpretation as protective opercula. We suggest that they served as opercula in addition to functioning as jaws. The primary function of the lower jaws was thus secondarily extended to that of protective shields when they acquired their calcitic covering, while as lower jaws their importance dwindled to that of a more passive abutment. Phylogenetically, this seems to have started slowly in some anaptychi and became obvious with the first aptychi. ▭ Ammonites, aptychus, operculum, jaw apparatus, evolution, function .     

New data on the jaw apparatus of fossil cephalopods

A newly discovered fossil cephalopod jaw apparatus that may belong to Permian representatives of the Endocochlia is described. Permorhynchus dentatus n. gen. n. sp. is established on the basis of this apparatus. The asymmetry of jaws in the Ectocochlia may be connected with the double function of the ventral jaw apparatus, and the well-developed, relatively large frontal plate of the ventral jaw should be regarded as a feature common to all representatives of ectocochlian cephalopods evolved from early Palaeozoic stock. Distinct features seen in the jaw apparatus of Upper Permian endocochlians include the pronounced beak form of both jaws and the presence of oblong wings on the ventral mandible.     

Über den Kieferapparat der Lytoceratacea (Ammonoidea)

The jaw apparati of Paleozoic and Triassic ammonoids are simìlar in shape to parrots’ beaks; they possess sharp cutting-edges and consist of chitinous material. Together with the similar jaw apparati of recent Coleoids, they constitute the Normal-Type of cephalopod jaws. The jaw apparati of the Jurassic-Cretaceous ammonites possess wide, shovel-like lower jaws without cutting-edges. They consist either of an undivided chitinous plate (Anaptychus) or of this and two additional calcitic plates (Aptychi) on its outer flanks. Jaw apparati of this type are here called Aptychus-Type jaws. New finds in the Upper Cretaceous of Japan suggest the existence of a third type of jaw apparatus which is here called the Rhynchaptychus-Type. Jaws of this type are mainly characterized by possession of calcitic rostra on the tips of the jaws. They seem to be restricted to members of the order Lytoceratacea (excluding the Heteromorphs).     

Mode of life and habitat of scaphitid ammonites

Scaphitid ammonites (scaphites) are among the most common ammonites in the Upper Cretaceous of the U.S. Western Interior. We have examined species of Hoploscaphites from the Campanian and Maastrichtian Pierre Shale and Bearpaw Shale for clues about their mode of life and habitat. Like most other ammonites, scaphites exhibit determinate growth. The “morphogenetic countdown” begins at the point at which the shell departs from the spiral coil and develops into a shaft and recurved hook. This is accompanied by a reduction in the spacing of ribs and tubercles on the hook and the formation of a constriction and varix at the aperture. Internally, this coincides with a decrease in septal spacing. The common association of scaphites and benthic fossils (e.g., Inoceramus) is interpreted as reflecting oxygen-rich episodes in the history of the Western Interior Seaway. Habitat depths are estimated at less than 100 m, based on faunal associations and studies of the mechanical strength of the septa and siphuncle. Analyses of the isotopic composition of the shells suggest that the animals lived near the sea floor. The high angle of orientation of the aperture at maturity (approximately 100°) seems incompatible with a nektobenthic mode of life. The constricted aperture ending in a thin lip may also have restricted unimpeded movement of long, muscular arms protruding from the aperture. Adults were probably poor swimmers, based on a comparison of their musculature (as inferred from their muscle scars) with that of nautilus (assuming that such a comparison is valid). The lack of a hyponomic sinus on the midventer would have prevented the animal from extending its hyponome below the shell in order to swim forward. As a consequence, scaphites may have been limited in their movement to swimming backward or downward, and may have exploited a low-energy lifestyle, remaining at a single site for an extended period of time. They may have consumed small prey in the water column, which is consistent with the presence of an aptychus-type lower jaw.     

Shell microstructure and mineralogy of the Littorinidae: ecological and evolutionary significance

An examination of the shell microstructure and mineralogy of species from 30 of the 32 genera and subgenera of the gastropod family Littorinidae shows that most species have a shell consisting of layers of aragonitic crossed-lamellar structure, with minor variations in some taxa. However, Pellilitorina, Risellopsis and most species of Littorina have partly or entirely calcitic shells. In Pellilitorina the shell is made entirely of calcitic crossed-foliated structure, while in the other two genera there is only an outer calcitic layer of irregular-prismatic structure. A cladistic analysis shows that the calcitic layers have been independently evolved in at least three clades. The calcite is found only in the outermost layers of the shell and in species inhabiting cooler waters of both northern and southern hemispheres. Calcium carbonate is more soluble in cold than warm water and, of the two polymorphs, calcite is about 35% less soluble than aragonite. We suggest that calcitic shell layers are an adaptation of high latitude littorinids to resist shell dissolution.     

Mosasaur predation on a nautiloid from the Maastrichtian Pierre Shale,Central Colorado,Western Interior Basin,United States

Mosasaurs were common predators on the ammonites that inhabited the upper water column of the Late Cretaceous Western Interior epicontinental seaway of North America. Mosasaurs developed predictable behaviour patterns for feeding on ammonite prey. There are no previous reports of mosasaur predation on the much less common Cretaceous nautiloids, possibly because of the prey's predominantly deep, epibenthic habitat, as deduced from modern Nautilus life habits. A single specimen of the highly inflated nautiloid, Eutrephoceras dekayi (Conrad), prey to a small adult mosasaur, likely Platycarpus, Prognathodon or Mosasaurus, is reported herein from the Pierre Shale of Colorado in the Early Maastrichtian biozone of Baculitesgrandis transitional to the biozone of B. clinolobatus. The nautiloid was killed in the same manner as described previously for discoid ammonites (Placenticeras, Sphenodiscus) from coeval strata in the USA and Canada.     

Aptychen als Kieferelemente der Ammoniten

The jaws of the ammonite generaEleganticeras, Hildoceras, Normannites, Scaphites, Physodoceras, andQuenstedtoceras (ofQuenst. lower jaws only) are described. Lower jaws and aptychi of these genera are shown to be identical. They consist of an inner layer of organic material, which corresponds to the complete anaptychi of liassic ammonites, and an outer layer of calcitic material deposited on both flancs of the lower jaw, giving rise, after decay of the organic layer, to the aptychi. Previously published interpretations of aptychi as opercula or hoods are discussed and rejected. It is concluded that they certainly functioned as jaws or possibly shovel-like devices.     

Paleoceanography of the Late Cretaceous (Maastrichtian) Western Interior Seaway of North America: evidence from Sr and O isotopes

Well-preserved fossils of the Late Cretaceous Western Interior Seaway (WIS) of North America have been analyzed for Sr concentration and Sr and O isotopes in order to decipher paleosalinities and paleotemperatures. The samples are from four biofacies within the Seaway (late Maastrichtian): offshore Interior (Pierre Shale), nearshore Interior (Fox Hills Formation), brackish (reduced salinity; Fox Hills Formation) and freshwater (Hell Creek Formation). Samples were also obtained from the Severn Formation of Maryland (considered to be representative of the open ocean). All biofacies (except the freshwater) are demonstrably within the Jeletzkytes nebrascensis ammonite zone (<1 Ma duration). The ⁸⁷Sr/⁸⁶Sr ratios show significant and systematic decreases from marine (mean±1 S.D.=0.707839±0.000024) to brackish facies (mean±1 S.D.=0.707677±0.000036), consistent with dilution by freshwater with a lower ⁸⁷Sr/⁸⁶Sr ratio than seawater. Such variation disallows using the ⁸⁷Sr/⁸⁶Sr ratios of fossil shell material to assign ages to fossils from the Late Cretaceous WIS without knowledge of the salinity in which the organism grew. The Sr isotope ratios for scaphitid ammonites within a single biofacies are similar to each other and different from those for scaphites in other biofacies, implying that these organisms are restricted in their distribution during life. The ⁸⁷Sr/⁸⁶Sr values of freshwater unionid mussels range widely and are not compatible with the freshwater endmember ⁸⁷Sr/⁸⁶Sr ratio required by the trend in ⁸⁷Sr/⁸⁶Sr vs. biofacies established from the other samples. Paleosalinities for the biofacies are estimated to range from 35‰ in the open marine to a minimum of 20‰ in the brackish, based on the presence of cephalopods in all four facies and the known salinity tolerance of modern cephalopods. Producing reasonable ⁸⁷Sr/⁸⁶Sr values for the freshwater endmember of a ⁸⁷Sr/⁸⁶Sr vs. 1/[Sr] plot requires a Sr concentration 0.2-0.5 that of seawater for the dominant freshwater input to the WIS. Such high Sr concentrations (relative to seawater) are not observed in modern rivers, and we suggest that the brackish environment in the WIS arose through the mixing of freshwater and seawater in a nearshore aquifer system. Reactions of the solution with aquifer solids in this ‘subterranean estuary’ [Moore, Mar. Chem. 65 (1999) 111-125] produced brackish water with the Sr concentration and isotopic composition recorded in the brackish biofacies. δ¹⁸O values of the fossils show decreases from the marine to brackish biofacies consistent with increasing temperatures (from ∼13 to 23°C) or, if temperatures were relatively constant, to a decrease in the δ¹⁸O of the water in which the shell formed. The latter interpretation is consistent with less-than-fully marine salinities in the nearshore biofacies, but both changes in temperature and the isotopic composition of the water may have occurred in this environment.     

Radula und Fangarme beiMichelinoceras sp. aus dem Silur von Bolivien

New finds of michelinoceratid cephalopods from the Kirusillas-Shale (Ludlow/Silurian) of Ushpa-Ushpa in the Eastern Cordillera of the Bolivian Andes reveal parts of the radula and imprints of arms owing to extremely good preservation conditions. The imprints of soft parts point that the orthoce-rates had 10 arms, two of them shaped to long tentacles as recent coleoids have. Of more importance is the first proof of a michelinoceratid radula in situ. This is the first known ra-dula of lower palaeozoic cephalopods at all. TheMichelinoceras radula consisted of 7 teeth per row. So the radulae of michelinoceratid cephalopods are very similar to those of ammonites and coleoids. But there are great differences in the radula ofMichelinoceras sp. and that ofNautilus sp. All known radulae of fossil and recent cephalopods are compared and phylogenetic or systematic implications are discussed. The classification of the cephalopods into six subclasses as used in the “Treatise” turns out to be unnatural and should be given up. Finally ecological relations between the morphology of radula-elements and mode of life are dis-cussed. Size and number of radula teeth let suppose that thisMichelinoceras specimen must have been an adult individual living far off the coast in pelagic seas.     

Skeletal chemistry of Nautilus and its taxonomic significance

The strontium (Sr) and magnesium (Mg) chemistry of the shell wall and septum as well as the spherulitic-prismatic and nacreous layers of these structures was determined for Nautilus species: N. belauensis, N. macromphalus, N. pompilius and N. scrobiculatus. Each species of Nautilus exhibits greater variability and higher concentrations of Mg in juvenile portions of the shell than in more mature portions of the shell. This decrease in the variability and amount of Mg in the aragonite lattice suggests a physiochemical system which becomes more efficient with time relative to carbonate production. Statistically significant differences in the Sr and Mg content of spherulitic-prismatic and nacreous layers of the shell and septum indicate that these layers were formed from extracellular fluids of different compositions. Concentrations of Sr and Mg in aragonite of the shell wall are characteristic for each species and sufficiently invariant within species to allow species of Nautilus to be distinguished statistically on the basis of either the Sr or Mg content of the shell wall or the Mg content of septa.     

Aptychi microstructure in Late Cretaceous Ancyloceratina (Ammonoidea)

The microstructure of aptychi (bivalved calcareous coverings on lower jaws) of three genera of Late Cretaceous Ancyloceratina, Baculites, Polyptychoceras and Jeletzkytes is described for the first time on the basis of well-preserved and in situ material from the Western Interior of the USA and Hokkaido, Japan. Optical and scanning electron microscope observations of aptychi on polished median and cross-sections reveal some variation in their relative size, shape and microstructure among the three genera. The aptychus of Baculites is composed of two calcitic layers: one with tilted lamellae and the other one with horizontal lamellae, whereas those of Polyptychoceras and Jeletzkytes consist of a thin layer with horizontal lamellae. Comparison with aptychi (e.g. Laevaptychus) of Jurassic Ammonitina shows that the aptychi of Ancyloceratina differ from those of Jurassic Ammonitina in the smaller number of layers and the absence of a sponge-like structure. We propose for the first time growth models for a sponge-like aptychus of Jurassic Ammonitina and the lamellar aptychus of Cretaceous Ancyloceratina. The remarkable microstructural variation of aptychi observed in Mesozoic Ammonoidea is probably related to the diversity of their modes of feeding and the secondary function of the lower jaws as opercula.     

Holography reveals the soft anatomy of ancient cephalopods

It is productive to view the suture-lines on fossil cephalopod shells as simple Fraunhoeffer diffraction patterns which can be inverted, by elementary Fourier Transformation, to provide a clear holographic image of the soft body-mass that produced them. As fleshy body organs are rarely ever preserved in ancient fossils it is particularly valuable to be able to deduce soft anatomy from conspicuous and usually well-preserved shell markings. Preliminary studies reported in this paper supply insight into the ontogeny and phylogeny of ancient cephalopods.     

Distribution of γ-carboxyglutamic acid in calcified tissues

γ-Carboxyglutamic acid, previously identified in the vertebrate mineralized tissues of bone and dentin, is not detectable in the calcified skeletons of six invertabrate species representing five phyla. Its absence in all analyzed invertebrate tissues (including calcitic, aragonitic, and apatitic mineral phases) indicates that matrix protein-bound γ-carboxyglutamic acid is not obligatory for the calcification process in the invertebrates. Further, these data raise the possibility that invertebrates as a group may lack the enzymatic capability for biosynthesizing γ-carboxyglutamic acid. In contrast, the distribution of γ-carboxyglutamic acid in the vertebrates has been further extended by this study to include an aptitic shark tooth and an aragonitic fish otolith. No γ-carnoxyglutamic acid was detected, however, in the organic matrix of the calcitic hen egg shell.     

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.     

Endothermic mosasaurs? Possible thermoregulation of Late Cretaceous mosasaurs (Reptilia,Squamata) indicated by stable oxygen isotopes in fossil bioapatite in comparison with coeval marine fish and pelagic seabirds

The thermoregulatory style of Late Cretaceous mosasaurs has become a highly controversial subject in vertebrate palaeontology. These extinct marine reptiles have previously been described as poikilothermic, endothermic or gigantothermic. Here we analyse three genera of mosasaurs from the Mooreville Chalk in Alabama (USA) of differing body mass, and compare their δ¹⁸O_PO4 derived body temperatures (T_b) with those of coeval poikilothermic fish (Enchodus) and endothermic pelagic seabirds (Ichthyornis). Results show that all mosasaurs, Clidastes (T_b = 33.1°C), Platecarpus (T_b = 36.3°C), and Tylosaurus (T_b = 34.3°C), had elevated average body temperatures in relation to those of the fish (T_b = 28.3°C) and were closer to those of Ichthyornis (T_b = 38.6°C). The temperatures calculated for Enchodus compare well with previously reported temperature estimates for the Mooreville Chalk and the T_b of Ichthyornis compares well with temperatures that have been reported for modern seabirds, suggesting that this method provides accurate results. Finally, although there are small differences of body temperature among mosasaur genera, these are independent of size, and thus inferred body mass, suggesting that mosasaurs were not gigantotherms, but rather endotherms.     

Recent advances in heteromorph ammonoid palaeobiology

Heteromorphs are ammonoids forming a conch with detached whorls (open coiling) or non-planispiral coiling. Such aberrant forms appeared convergently four times within this extinct group of cephalopods. Since Wiedmann's seminal paper in this journal, the palaeobiology of heteromorphs has advanced substantially. Combining direct evidence from their fossil record, indirect insights from phylogenetic bracketing, and physical as well as virtual models, we reach an improved understanding of heteromorph ammonoid palaeobiology. Their anatomy, buoyancy, locomotion, predators, diet, palaeoecology, and extinction are discussed. Based on phylogenetic bracketing with nautiloids and coleoids, heteromorphs like other ammonoids had 10 arms, a well-developed brain, lens eyes, a buccal mass with a radula and a smaller upper as well as a larger lower jaw, and ammonia in their soft tissue. Heteromorphs likely lacked arm suckers, hooks, tentacles, a hood, and an ink sac. All Cretaceous heteromorphs share an aptychus-type lower jaw with a lamellar calcitic covering. Differences in radular tooth morphology and size in heteromorphs suggest a microphagous diet. Stomach contents of heteromorphs comprise planktic crustaceans, gastropods, and crinoids, suggesting a zooplanktic diet. Forms with a U-shaped body chamber (ancylocone) are regarded as suspension feeders, whereas orthoconic forms additionally might have consumed benthic prey. Heteromorphs could achieve near-neutral buoyancy regardless of conch shape or ontogeny. Orthoconic heteromorphs likely had a vertical orientation, whereas ancylocone heteromorphs had a near-horizontal aperture pointing upwards. Heteromorphs with a U-shaped body chamber are more stable hydrodynamically than modern Nautilus and were unable substantially to modify their orientation by active locomotion, i.e. they had no or limited access to benthic prey at adulthood. Pathologies reported for heteromorphs were likely inflicted by crustaceans, fish, marine reptiles, and other cephalopods. Pathologies on Ptychoceras corroborates an external shell and rejects the endocochleate hypothesis. Devonian, Triassic, and Jurassic heteromorphs had a preference for deep-subtidal to offshore facies but are rare in shallow-subtidal, slope, and bathyal facies. Early Cretaceous heteromorphs preferred deep-subtidal to bathyal facies. Late Cretaceous heteromorphs are common in shallow-subtidal to offshore facies. Oxygen isotope data suggest rapid growth and a demersal habitat for adult Discoscaphites and Baculites. A benthic embryonic stage, planktic hatchlings, and a habitat change after one whorl is proposed for Hoploscaphites. Carbon isotope data indicate that some Baculites lived throughout their lives at cold seeps. Adaptation to a planktic life habit potentially drove selection towards smaller hatchlings, implying high fecundity and an ecological role of the hatchlings as micro- and mesoplankton. The Chicxulub impact at the Cretaceous/Paleogene (K/Pg) boundary 66 million years ago is the likely trigger for the extinction of ammonoids. Ammonoids likely persisted after this event for 40–500 thousand years and are exclusively represented by heteromorphs. The ammonoid extinction is linked to their small hatchling sizes, planktotrophic diets, and higher metabolic rates than in nautilids, which survived the K/Pg mass extinction event.