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 .     

Preferential predatory peeling: Ammonoid vs. nautiloid shells from the Upper Carboniferous of Texas,USA

Narrow groove-like excavations on ammonoid and coiled nautiloid shells are rare in Upper Carboniferous units from Texas, USA. The morphological characteristics of the excavation grooves typically are confined to the ventral and ventrolateral parts of the outer whorl of the shell, are narrower than the length, and have irregular edges where small segments or chips of shells have been removed. Analysis of these features reveals a statistically significant preferential occurrence on ammonoids (1.195% of ca. 3515 specimens) as compared to coiled nautiloids (0.506% of ca. 2965 specimens). The ammonoids typically have longer excavations that penetrate the phragmocone more frequently than those observed in the coiled nautiloids. The groove-like excavations were probably formed by the removal and peeling of shell material by one or more predatory or scavenging arthropods to obtain organic material (tissue and membranes) within the ammonoid and nautiloid body chambers and phragmocones. The excavations probably occurred when the cephalopod was alive (i.e., the cause of death) or shortly after the cephalopod's death. There is no evidence that the excavations are related to sheltering by the excavating organism.     

Origin of intracameral sheets in ammonoids

The distribution, morphology and mutual relationships of cameral sheets in ammonoids are revised and re-evaluated. Taking into account recent models of ammonoid septum and chamber formation, three different origins can be attributed to the morphological types of sheets: (1) membranes replicated by the rear mantle (pseudosepta and septal linings), (2) membranes secreted sequentially and/or stretched across the chamber (horizontal membranes and chamber linings) and (3) products of desiccation of the cameral liquid (transverse and siphuncular sheets), presumably a cameral hydrogel. Sheets are always preserved near the siphuncular area, because as the cameral liquid was pumped out from the chamber it became progressively richer in dissolved mucus. In the last-formed drops, or menisci, this mucus adhered to the surface of the previously secreted sheets and, on dehydration, it also replicated the surface of the residual reservoirs, producing desiccation sheets. On the basis of the new evidence, changes in the shape of the rear mantle in Triassic ammonoids can be reconstructed. In general, deformations affected the rounded or bottle-neck saddles, which deflated after detachment of the last-formed septum and reinflated when the position of the next septum was reached. The rest of the elements of the septal epithelium were affected to a much lesser extent. One of the functions of those cameral sheets secreted by the rear body was related to a more efficient transport of the cameral liquid upon decoupling from the siphuncular tube. Ammonoids, Triassic, septum, chamber growth, cameral sheets.     

Cameral membranes in prolecanitid and goniatitid ammonoids from the Permian Arcturus Formation, Nevada, USA

We describe cameral membranes in prolecanitid and goniatitid ammonoids from the Lower Permian Arcturus Formation, Nevada, USA. The membranes are preserved as phosphatic sheets and were originally composed of organic material such as conchiolin. Because the phragmocones are filled with micritic calcite, the cameral membranes can be exposed by etching with weak acetic acid. The membranes are associated with the siphuncle and also coat the septal faces and chamber walls. The siphuncular membranes are much more extensive in the prolecanitids than in the goniatites. These membranes appear in the prolecanitids at the beginning of the third whorl, corresponding to a shell diameter of 3-4 mm, and become more complex through ontogeny. Additional membranes, called transverse membranes, appear in some of the septal saddles on the ventrolateral side. The siphuncular membranes in prolecanitids are very similar to those in the Ceratitina plus Mesozoic Ammonoidea, suggesting that such membranes are widely distributed in this group. However, the origin and function of these membranes are unclear. We argue that the siphuncular membranes were sequentially secreted by the rear mantle during forward movement of the body and were not produced by desiccation of cameral liquid after the formation of the chambers. The most compelling arguments for this interpretation are the abrupt appearance of these membranes at a shell diameter of approximately 3-4 mm in prolecanitids, ceratites, and ammonitids, coincident with the end of the neanic stage, and the uniform increase in complexity of the membranes through ontogeny. The shape of the siphuncular membranes in prolecanitids suggests the presence of an invagination on the dorsal side of the siphuncle during part of the chamber formation cycle. Cameral membranes may have served a variety of functions including stabilizing the cameral liquid to reduce rocking motion during swimming, anchoring the siphuncle to the chamber wall, and facilitating cameral liquid removal, permitting a faster rate of growth.     

Buoyancy of some Palaeozoic ammonoids and their hydrostatic properties based on empirical 3D‐models

The interpretation of the function of the ammonoid phragmocone as a buoyancy device is now widely accepted among ammonoid researchers. During the 20^th century, several theoretical models were proposed for the role of the chambered shell (phragmocone); accordingly, the phragmocone had hydrostatic properties, which enabled it to attain neutral buoyancy, presuming it was partially filled with gas. With new three‐dimensional reconstructions of ammonoid shells, we are now able to test these hypothetical models using empirical volume data of actual ammonoid shells. We investigated three Palaeozoic ammonoids (Devonian and Carboniferous), namely Fidelites clariondi, Diallagites lenticulifer and Goniatites multiliratus, to reconstruct their hydrostatic properties, their syn vivo shell orientation and their buoyancy. According to our models, measurements and calculations, these specimens had aperture orientations of 19°, 64° and 125° during their lives. Although none of our results coincide with the aperture orientation of the living Nautilus, they do verify the predictions for shell orientations based on published theoretical models. Our calculations also show that the shorter the body chamber, the poorer was the hydrodynamic stability of the animal. This finding corroborates the results of theoretical models from the 1990s. With these results, which are based on actual specimens, we favour the rejection of hypotheses suggesting a purely benthonic mode of life of ammonoids. Additionally, it is now possible to assess hydrodynamic properties of the shells through ontogeny and phylogeny, leading to insights to validate theoretical modes of life and habitat through the animal's life.     

Empirical 3D model of the conch of the Middle Jurassic ammonite microconch Normannites: its buoyancy,the physical effects of its mature modifications and speculations on their function

A 3D model of the Middle Jurassic ammonoid Normannites with an apertural modification from Thürnen, Switzerland, was constructed using physical–optical tomography. It was tested to determine whether the formation of the apertural modification affected shell orientation, to estimate buoyancy regulation and to reconstruct the mode of life of this ammonoid. No drastic postural changes occurred between the 3D models that excluded and included lappets, suggesting that the lappets were not formed to change the syn vivo shell orientation and, in turn, locomotion. We speculate that these adult shell modifications served to protect the soft parts during the reproduction period. Buoyancy calculations based on the model assume that ammonoids were positively buoyant when the phragmocone was devoid of liquid. When 31% of the entire phragmocone was filled with liquid, the living animal would have reached neutral buoyancy in contrast to 27% of cameral liquid filling when the weight of the aptychi is included. Provided that smaller ammonoids had more cameral liquid than bigger ammonoids, such as the modern Nautilus, Normannites examined in this study would have been able to maintain neutral buoyancy and might have had a demersal, nektobenthic or nektonic habitat somewhere in the water column.     

Ventral bite marks in Mesozoic ammonoids

Reports on the predators of ammonoids are rare, although ammonoids were abundant and diverse invertebrates in many Paleozoic and Mesozoic marine ecosystems. Most previous work on lethal ammonoid predation has focused on (sub)circular tooth marks which resulted from fish and mosasaur attacks. In the present study we discuss a relatively common type of bite mark in ammonoid shells, the ‘ventral bite mark’. This typically occurs in a restricted position on the ventral side of the outer body chamber whorl and does not affect either the aperture or the phragmocone. Ammonoid specimens revealing ventral bite marks used in this study were collected from a wide range of strata which range in age from the Lower Jurassic to the uppermost Cretaceous (close to the Cretaceous–Paleogene boundary). These ventral bite marks are absent in the Paleozoic collections studied. The vast majority of ventral bite marks are situated at the end of the body chamber, close to the phragmocone. This is interpreted as the result of predatory attacks on the back or blind side of ammonoids in their living position. The predators aimed for the vital parts and muscle attachments to obtain the edible soft tissues. The agents for most of the ventral bite marks to ammonoids are probably coleoid cephalopods (especially teuthoids) and predatory fishes to a lesser extent.     

The Evolution and Development of Cephalopod Chambers and Their Shape

The Ammonoidea is a group of extinct cephalopods ideal to study evolution through deep time. The evolution of the planispiral shell and complexly folded septa in ammonoids has been thought to have increased the functional surface area of the chambers permitting enhanced metabolic functions such as: chamber emptying, rate of mineralization and increased growth rates throughout ontogeny. Using nano-computed tomography and synchrotron radiation based micro-computed tomography, we present the first study of ontogenetic changes in surface area to volume ratios in the phragmocone chambers of several phylogenetically distant ammonoids and extant cephalopods. Contrary to the initial hypothesis, ammonoids do not possess a persistently high relative chamber surface area. Instead, the functional surface area of the chambers is higher in earliest ontogeny when compared to Spirula spirula. The higher the functional surface area the quicker the potential emptying rate of the chamber; quicker chamber emptying rates would theoretically permit faster growth. This is supported by the persistently higher siphuncular surface area to chamber volume ratio we collected for the ammonite Amauroceras sp. compared to either S. spirula or nautilids. We demonstrate that the curvature of the surface of the chamber increases with greater septal complexity increasing the potential refilling rates. We further show a unique relationship between ammonoid chamber shape and size that does not exist in S. spirula or nautilids. This view of chamber function also has implications for the evolution of the internal shell of coleoids, relating this event to the decoupling of soft-body growth and shell growth.     

Taphonomic differentiation of Oxfordian ammonites from the Cracow Upland, Poland

Taphonomic analysis of Lower and Middle Oxfordian ammonites from the Cracow Upland, southern Poland (localities at Pod???e, Zalas, M?ynka) revealed differences in ammonite preservation. The studied ammonites, usually termed as external and internal moulds, show a more complex state of preservation. In the Middle Oxfordian glauconitic marls, ammonites are preserved as internal moulds with neomorphic calcite shells showing relics of the original internal structure. In the Middle Oxfordian platy peloidal limestones, ammonites are preserved mostly as external moulds, without septal suture, however under microscope might show relics of internal whorls and septa and/or subtle differences in sediment filling phragmocone chambers. In sponge–microbial bioherms and biostromes, ammonite internal moulds have shells, which in contrast to ammonites from glauconitic marls are not strictly neomorphic ones, but originated by shell dissolution and subsequent filling of moldic porosity by calcite cement. In sponge–microbial nodular limestones, the ammonites are strongly deformed and the outer wall is usually removed by dissolution under pressure. Other important taphonomic differences include the rate of compaction (highest in platy limestones), sedimentary infillings, microborings, encrustations and preservation of siphuncular tubes. The majority of the ammonites appear to be phragmocones; aptychi in all facies are rare. Siphuncular tubes are fossilized exclusively in oppeliids, only in specimens from glauconitic marls and platy limestones, although their other taphonomic attributes are different. Tubes seem to have fossilized due to microbially mediated phosphatization that could be favoured by a set of parameters which operated rather at the scale of ammonoid carcasses: closed, poorly oxygenated conditions, and reduced pH. Taphonomic processes were controlled by the sedimentary environment (fragmentation, sedimentary filling, phosphatization of siphuncular tubes), as well as by early and late diagenesis (neomorphic transformation, dissolution, cementation, compaction) influenced by lithology.     

Morphogenetic significance of the conchal furrow in nautiloids: evidence from early embryonic shell development of Jurassic Nautilida

As reported by many workers over the past two centuries, the inner part of the shell of various straight and coiled Palaeozoic to tertiary nautiloid taxa bears a continuous mid-ventral furrow that extends into the phragmocone and the body chamber nearly to the aperture. Study of the early embryonic shell development of Jurassic Nautilida shows that the most apical part of this so-called conchal furrow originates from the inner part of the initial, calcified shell apex, in line with the inner ventral termination of the central linear depression of the cicatrix, the initial site of shell deposition. The conchal furrow corresponds to a morphological feature arising as a developmental by-product. Rare specimens of scattered ammonoid species (and possibly of bactritoids) display a similar feature, whereas their protoconch lacks a cicatrix. However, the protoconch of recent cuttlefish, Sepia officinalis, often displays a longitudinal fold of the primary shell epithelium. A longitudinal groove or a pair of grooves appears connected with this cicatrix-like structure. Although the mid-ventral ridge in ammonoids must probably be viewed as an incidental 'fabricational noise', whether or not it originates from a so far undocumented optional ridge on the protoconch or from some other structure related to shell development remains an open question.     

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 .     

Composition and ontogeny ofDictyoconites (aulacocerida,cephalopoda)

Dictyoconites from the middle Triassic Cassian Formation is a characteristic representative of the Aulacocerida. Embryonic development and construction of the phragmocone is like that of Jurassic belemnites. The siphuncular tube is double-walled with a long retrochoanitic mineralized septal neck continuing into an organic tube. The extended decoupling zone resembles that ofSpirula. Characteristic ofDictyoconites are the tubular »living chamber« and two layered deposits of the muscular mantle on the phragmocone. The Triassic coleoid was a slender squid with visceral mass and mantle cavity encapsuled in shell and the whole conch covered by muscular mantle extending in two lateral apical fins attached to the aragonitic rostrum.     

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.     

Rhythms of ammonoid shell secretion

From measurements of the thickness of successive growth rings shell growth curves were constructed for seven ammonoid genera of different taxonomic position and geological age. All the curves have 'sinusoidal' aspect due to the alternation of maximum and minimum growth increments of the shell tunc. Growth curves made for two ammonoid shells belonging to the genera Proloxyclymenia (D₃fm) and Euphylloceras (K₁a) show regularly recurring minima and maxima at intervals of 14–16 growth rings. The fact that the minima and maxima of the growth curves recur exactly at intervals of 14–16 growth rings seems to reflect the relationship between the shell tube secretion and the formation of septa. This fact also seems to support the views about fortnightly or 28-day cycles in the construction of ammonoid hydrostatic chambers. By counting the total number of septa in the phragmocones, the duration of phragmocone secretion for 23 shells was estimated. Preliminary inferences on the presence in ammonoid shells of daily, bidaily and weekly growth rings are made. □ Ammonoidea, growth rhythms.     

Irregular calcareous sheets preserved in a conch of the Cretaceous ammonoid Hypophylloceras from Hokkaido,Japan

The mantle tissue is essential for understanding the diverse ecology and shell morphology of ammonoid cephalopods. Here, we report on irregular calcareous sheets in a well-preserved shell of a Late Cretaceous phylloceratid ammonoid Hypophylloceras subramosum from Hokkaido, Japan, and their significance for repairing the conch through the mantle inside the body chamber. The sheets are composed of nacreous layers arranged parallel to the irregularly distorted outer whorl surface. The nacreous sheets formed earlier are unevenly distributed and attached to the outer shell wall locally, whereas the last formed sheet covers a wide area of the outer shell wall. The absence of any interruption of ribbing around the irregular area suggests that these sheets were secreted inside the body chamber from the inner mantle. Gross morphological and X-ray computed tomography observations revealed that the spacing of septal formation was not affected by this event. The complex structure of the irregular sheets suggests a highly flexible mantle inside the body chamber.     

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.     

Untangling phylogenetic,geometric and ornamental imprints on Early Triassic ammonoid biogeography: a similarity‐distance decay study

Ammonoids are diverse and widespread fossil, externally shelled cephalopods that flourished for more than 300 Myr before their total extinction 65 Ma ago. In spite of two centuries of intensive scientific studies, their mode(s) of life and long‐distance dispersal abilities remain poorly known. Here, we address this by focusing on the latitudinal distribution of Early Triassic (approximately 250 Myr) ammonoids through similarity‐distance decay analyses. We examine and compare rates of similarity‐distance decay between various groups with respect to systematics, shell geometry and ornamentation to untangle phylogenetic, geometric and ornamental imprints on the observed biogeographical pattern. Our data do not support any phylogenetic and shell ornamentation influence, but rather demonstrate the significant effect of (sub‐)adult shell geometry on the similarity–distance decay: most evolute morphs tend to have been more endemic than most involute forms. This contrasts with the classic hypothesis that long‐distance ammonoid dispersal mainly occurred during the earliest planktonic stages, and thus that (sub‐)adult morphological characteristics should not constrain large‐scale biogeographical patterns of ammonoids. Although direct control by Sea Surface Temperature can be discarded, this result may indicate that at least some adult Triassic ammonoid morphs were skilled active swimmers capable of achieving long‐distance migration, as observed for some present‐day coleoid cephalopods. □Ammonoid, dispersal, similarity‐distance decay, morphology, phylogeny, biogeography, Triassic.     

The role of suture complexity in diminishing strain and stress in ammonoid phragmocones

Several hypotheses have been put forward to explain the sinuosity and complexity of suture lines in Ammonoidea. At present, the two principal opponent views maintain either that high complexity was a requisite to reinforce the shell in response to hydrostatic pressure, or that complexity augmented the attachment area for muscles. By using finite element calculations and analytical estimates of simplified ammonoid shell geometries, it is shown that complex suture lines reduced dramatically the strain and the stress in the phragmocone. The calculations lend support to the hypothesis that high sinuosity is an evolutionary response to external pressure. Additionally, it is found that without complex septa, the inward deformation of an ammonoid with thin shell would cause it to shrink in response to pressure and to lose buoyancy by a non-negligible amount.     

The black layer in cephalopods from the German Muschelkalk (Triassic)

Thin, dark, probably phosphatic coatings were found on the dorsum in front of and sometimes behind the aperture of 50 specimens of Paraceratites and Ceratites (Ammonoidea) belonging to 14 species and subspecies and in three specimens of Germanonautilus , all from the Middle Triassic of Germany. The proportions, occurrences, position, outline, and preservation in fossil Nautiloidea and Ammonoidea (originally organic matter) of this structure support the hypothesis that it is homologous with the black layer in Recent Nautilus and Allonautilus . It is not yet possible to test whether these cephalopods show homologous styles of the development of these structures or whether the black layer can be identified in a common ancestor. In contrast to many ammonoids, Ceratites and Paraceratites , most Palaeozoic ammonoids, and some Mesozoic ammonoids probably did not have lower mandibles that were suitable for the closure of the aperture. They probably possessed a dorsally extending mantle (supracephalic mantle fold) and a hood, as in Recent Nautilus and Allonautilus , that was attached to the black layer. This interpretation is corroborated by a similar morphology of the black layer in an adult specimen of the nautilid Cenoceras from the South German Middle Jurassic and three specimens of Germanonautilus from the South German Middle Triassic (both Nautiloidea).