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.     

Early evolutionary trends in ammonoid embryonic development

During the Devonian Nekton Revolution, ammonoids show a progressive coiling of their shell just like many other pelagic mollusk groups. These now extinct, externally shelled cephalopods derived from bactritoid cephalopods with a straight shell in the Early Devonian. During the Devonian, evolutionary trends toward tighter coiling and a size reduction occurred in ammonoid embryonic shells. In at least three lineages, descendants with a closed umbilicus evolved convergently from forms with an opening in the first whorl (umbilical window). Other lineages having representatives with open umbilici became extinct around important Devonian events whereas only those with more tightly coiled embryonic shells survived. This change was accompanied by an evolutionary trend in shape of the initial chamber, but no clear trend in its size. The fact that several ammonoid lineages independently reduced and closed the umbilical window more or less synchronously indicates that common driving factors were involved. A trend in size decrease of the embryos as well as the concurrent increase in adult size in some lineages likely reflects a fundamental change in reproductive strategies toward a higher fecundity early in the evolutionary history of ammonoids. This might have played an important role in their subsequent success as well as in their demise.     

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 .     

Life-cycles of some Devonian ammonoids

Whorl expansion rates of six representative ammonoid genera from late Emsian and Eifelian strata of Morocco were calculated for each whorl. The corresponding body chamber lengths and the orientations of the apertures were computed based on these values. The resulting body chamber length and orientation of the aperture graphs were compared with other conch features, ecology of Recent cephalopods, and sedimentological data of the host rocks in the Tafilalt (eastern Anti-Atlas, Morocco). A subdivision of the ontogeny of these ammonoids was achieved comprising the early and late embryonic periods, the juvenile period, the preadult, and the adult growth period. All growth periods are defined by specific changes in growth, conch morphology, and mode of life. According to this reconstruction, hatchlings were probably already capable of active movements. Differentiation in two main modes of life of the examined taxa occurred in the late juvenile or early preadult period. As preadult animals, most of the Mimagoniatitoidea and Agoniatitoidea became active swimmers (Nektonic), whereas the representatives of the Anarcestoidea were capable of slow movements only (Planktonic). As adults, most representatives of the three superfamilies had an approximately horizontally oriented aperture, allowing active swimming and possibly active choice of spawning sites. Additionally, the new ammonoid taxon Rherisites tuba gen. nov., sp. nov. from the late Emsian is introduced.     

Covariation of morphological characters in the Triassic ammonoid Czekanowskites rieberi

The Triassic ammonoid Czekanowskites rieberi displays a covariation of morphological charac ters, which is rather common in ammonoids. Its morphological spectrum ranges from laterally compressed, involute, weakly ribbed forms to depressed, semiinvolute, strongly ribbed forms. In order to study this covariation, fifteen axially cut specimens have been analyzed by means of image analysis, which allows us to obtain the ontogenetic record of radii, area and perimeter of the individual whorl cross-sections. A logarithmic model of growth has been applied. Our data indicate that, owing to the covariation, the radii from the origin to the venter and to the umbil ical seam of a given whorl section vary inversely in order to maintain the relative position of the center of gravity of the whorl cross-section both throughout the ontogeny of single specimens and within the population. This influences hydrostatic parameters, such as the position of the center of mass and the orientation and stability of the shell. Since the ontogenetic record of the angular length of the body chamber is not known, we have calculated those hydrostatic varia bles using two mutually exclusive assumptions: (1) the angular length of the body chamber was constant throughout ontogeny and (2) the volume of the body chamber grew monotonically with the revolution angle. Fluctuations of the three hydrostatic variables were always less important in the first assumption. In any case, the spectrum of, for example, theoretical orien tations is comparable to those observed in the species of present-day Nautilus. The range of adult body-chamber length observed in C. rieberi is much narrower than the theoretical adult body-chamber length calculated under the second assumption which indicates that a certain control over this parameter existed in the natural population, probably in order to maintain a narrow range in orientation and stability. The excess or deficit in soft-body weight was probably compensated by inverse variations in shell-wall weight. The main conclusion is that, despite the extreme morphological variability, hydrostatic and, possibly, hydrodynamic properties of the population remained within narrow limits.     

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.     

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.     

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.     

Allometric growth and intraspecific variability in the basal Carboniferous ammonoid<Emphasis Type="Italic">Gattendorfia crassa</Emphasis><Emphasis Type="SmallCaps">Schmidt</Emphasis>, 1924

Gattendorfia crassa is an Early Carboniferous (Mississippian) goniatite species with strikingly allometric conch growth. Analysis of 15 high-precision cross-sections of this species demonstrates the small intraspecific variability of some of the conch form characters, but remarkable variability in others. While the whorl expansion rate, umbilical width, and conch thickness vary within narrow limits, the expansion rates of the whorl height and whorl width are remarkably plastic. Variability of most of the characters tends to be smallest in intermediate growth stages, whereas juveniles and adults are more variable. The differences in morphological plasticity are interpreted in terms of the function of the ammonoid conch, especially the orientation of the aperture during life.      

From near extinction to recovery: Late Triassic to Middle Jurassic ammonoid shell geometry

The Triassic–Jurassic extinction resulted in the near demise of the ammonoids. Based on a survey of ammonoid expansion rates, coiling geometry and whorl shape, we use the Raup accretionary growth model to outline a universal morphospace for planispiral shell geometry. We explore the occupation of that planispiral morphospace in terms of both breadth and density of occupation in addition to separately reviewing the occurrence of heteromorphs. Four intervals are recognized: pre‐extinction (Carnian to Rhaetian); aftermath (Hettangian); post‐extinction (Sinemurian to Aalenian) and recovery (Bajocian to Callovian). The pre‐extinction and recovery intervals show maximum disparity. The aftermath is marked by the disappearance of heteromorphs and a dramatic reduction in the range of planispiral morphologies to a core area of the morphospace. It is also characterized by an expansion into an evolute, slowly expanding part of the morphospace that was not occupied prior to the extinction and is soon abandoned during the post‐extinction interval. Aftermath and post‐extinction ammonoid data show a persistent negative correlation whereby rapid expansion rates are associated with narrow umbilical widths and often compressed whorls. The permanently occupied core area of planispiral morphospace represents generalist demersals whose shells were probably optimizing both hydrodynamic efficiency and shell stability. All other parts of the planispiral morphospace, and the pelagic modes of life the shells probably exploited, were gradually reoccupied during the post‐extinction interval. Planispiral adaptation was by diffusion away from the morphospace core rather than by radical jumps. Recovery of disparity was not achieved until some 30 Myr after the extinction event.     

Morphospace occupation of ammonoids over the Devonian-Carboniferous boundary

In a case study, a stratophenetic analysis of the succeeding ammonoid faunas of the latest Devonian and earliest Carboniferous cephalopod limestones of the Rhenish Massif has been made. This investigation concentrated on the development of the whorl expansion rate (WER), a character very important for ammonoids since it indicates the body chamber length and hence the orientation in the water column as well as mobility. The study leads to the conclusion that the Hangenberg Event caused an almost complete change in the morphospace adopted by ammonoids. All clymenüds as well as tornoceratids became extinct at or immediately after the Hangenberg Event, and the morphospace left behind by these Devonian groups was reoccupied only incompletely by the surviving prionoceratid ammonoids.     

<Emphasis Type="Italic">Goniatites sphaericus</Emphasis> (Sowerby, 1814), the archetype of Palaeozoic ammonoids: a case of decreasing phenotypic variation through ontogeny

The Late Viséan (Early Carboniferous) ammonoid species Goniatites sphaericus (Sowerby, 1814) is revised with the use of the type material. With respect to conch morphology, suture line and particularly shell ornament, it is regarded as a senior synonym of the species Goniatites fimbriatus (Foord and Crick, 1897) and must therefore be re-established as an important index fossil for the ammonoid stratigraphy of the Rhenohercynian and Subvariscan shelves. The Goniatites fimbriatus Zone has been re-named Goniatites sphaericus Zone on this basis. The study of ontogenetic changes in the conch parameters shows that intraspecific variation occurs within a wide range. In the whorl profile, variation is very high in small juveniles, while in the coiling rate, intermediate growth stages are more strongly affected.     

Morphological and structural adaptations to soft substrates in the Early Jurassic monomyarians Lithiotis and Cochlearites

Lithiotis problematica and Cochlearites loppianus are sessile monomyarian bivalves known from the Early Jurassic of the Tethyan region, where they are found standing vertically in the calcareous mud of lagoonal fades. Their shells are characterized by a long cardinal area, ventrally displaced body space, thick attached valve, and a thin, flat free valve of equal length. A functional ligament is present only in juvenile stages of Cochlearites. To grow straight, the ventral end of the shell had to gape, so in both species it was probably the elasticity of the thin free valve that caused the shell to open. In the adult stage the shell grew only towards the venter with an apparently constant growth rate, while the soft body size remained unchanged. The outer shell is composed of a compact aragonitic layer, while the inner part is filled with loose chalky deposits which are thought to have functioned as supporting the soft body and lifting it upwards. These and other morphologic and structural features, as well as the growth pattern, can be explained as an adaptation of a sessile animal to soft muddy bottoms and rapid sedimentation. Some elongated Crassostrea living in similar environments show remarkable morphologic and structural convergence with Lithiotis and Cochlearites.     

Ammonoid palaeobiology

This, the second in the series of virtual issues of Palaeontology, charts the development of concepts in ammonoid palaeobiology, with reference to nine selected papers published in this journal from 1965 onwards. These cover a broad range of topics: goniatite bed palaeoecology, systematics and evolution in Scaphites, flow dynamics of cephalopod shells, shell growth and differential geometry, septal function, ammonoid assemblage analysis, evolutionary trends, stratocladistics, and soft part preservation. Despite the growth and success of palaeobiological interpretations of ammonoid evolution, life, morphology and environments over the last 50 years, ammonoid research retains a great potential for the future.     

Selective settlement of the barnacle Semibalanus balanoides (L.) facilitates its growth and reproduction on mussel beds in the Wadden Sea

On the unstable sedimentary tidal flats of the Wadden Sea, a suitable attachment substrate for sessile organisms is generally lacking. Epibenthic mussel beds (Mytilus edulis L.) provide the only and strongly limited settlement sites available for the barnacle, Semibalanus balanoides (L.). Field investigations showed that barnacles were non-randomly distributed within a mussel bed. They preferentially occurred near the siphonal apertures of living mussels but rarely grew on dead mussels or shell fragments. Field experiments revealed that this was due to selective settlement of barnacle cyprid larvae. Growth of barnacles was significantly higher upon living mussels than on empty mussel shells. Moreover, a higher reproductive output was obtained by individuals on living mussels which produced twice as many nauplii larvae than barnacles attached to empty shells. This study shows that selective settlement of S. balanoides cyprid larvae on living mussels is adaptive with respect to individual fitness. Received in revised form: 15 January 2001 Electronic Publication     

Shell morphology and microstructure of Neogene <Emphasis Type="Italic">Melanopsis</Emphasis> Ferussac (Gastropoda)

Representatives of the genus Melanopsis have an extraconch, a shell structure unusual for gastropods, which envelops several whorls, including the last whorl and up to two preceding whorls. Microstructural features of the shell are studied for the first time, using M. impressa Krauss as an example. It is shown that the extraconch is formed by the outer crossed-lamellar layer.     

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.     

Buckman's Paradox: variability and constraints on ammonoid ornament and shell shape

Buckman's Law of Covariation states that ammonoid shell shape and ornamentation are typically correlated, such that compressed, involute forms have light ornament while more inflated, evolute forms have heavier ornament. Such covariation has been observed in many ammonoid groups, and implies a link between the morphogenesis of shell shape and ornamentation. However, other evidence suggests that while ornament growth is controlled by the genetic‐developmental program of the ammonoid, shell shape is strongly influenced by environmental factors. These differing viewpoints lead to Buckman's Paradox – are ornamentation and shell shape tightly linked, as implied by Buckman's covariation, or is the morphogenesis of ornament controlled genetically, while shell shape is controlled environmentally? To address this issue, the variability of shell shape and rib morphology has been compared for a group of endemic acanthoceratid ammonites from the Cretaceous Western Interior Seaway of North America. If Buckman's Law holds due to a morphogenetic connection between shell shape and ornamentation, we would expect taxa with more variable shell shapes to also show more variable rib features and growth. Morphometric analysis of seven shell shape and two rib characters for the Western Interior acanthoceratids finds no such correlation, suggesting that shell shape and rib growth are controlled by different processes. Indeed, rib growth appears to be more constrained than shell shape, consistent with the view that ornamentation is more tightly controlled by the developmental‐genetic growth program of the ammonoid. These results emphasize the complexity of ammonoid morphogenesis and highlight our limited understanding of the causes underlying Buckman's Law.     

A paradox survival – report of a repaired syn vivo perforation in a nautiloid phragmocone

The nautiloid Trocholites depressus (Eichwald, 1840) from the Lasnamägi regional stage (Darrivillian, Middle Ordovician) of Vaike Pakri Island (North-West Estonia) is the only known ectocochleate cephalopod that survived and healed a perforation of the phragmocone. Two chambers of the specimen were broken during its lifetime. The injury is located on the venter of the conch directly above the peristomal opening of the body chamber. It is reconstructed that the peristomal mantle tissue carried out an initial sealing of the injured chambers. The complete calcified sealing and compensation of the irregular shell surface started late with the overgrowth of the septa of the preceding whorl. The position and diameter of the siphuncle were not disturbed by the regeneration showing that these characters allowed a low phenotypic variability. Despite the trauma in the midlife growth history of this specimen, it appears to have reached maturity.