Limpet home depressions in Cretaceous ammonites

Remarkable large pits were found on the shell surface of many ammonites from the Turonian-Maastrichtian sediments of Hokkaido, northern Japan, and Sakhalin, Russia. These pits are (1) round to elliptical in shape; (2) up to 20 mm (usually around 10 mm) in diameter; (3) shallow depressions to deep holes that almost penetrate the shell: (4) occasionally healed by a thin shell blister from inside the shell; (5) often overlapping one another; (6) several to more than 170 in number on one flank of the ammonites; (7) found on both flanks and predominantly on the body chamber and the final volution of the phragmocone; and (8) found only in the two families Pachydiscidae and Puzosiidae, predominantly of more than 300 mm in shell diameter. They can best be interpreted as the home depressions of patellogastropod limpets. The presence of pits on both flanks of the ammonites and those healed from inside the shell strongly suggest that the limpets were dwelling on mature swimming ammonites. Host specifity, their very small shell size compared with the host ammonites, and sparse Occurrence in sediments favor a mode of life as obligate pseudoplankton. We suggest that this remarkable limpet-ammonite association was well established in northwestern Pacific bioprovinces during the late Cretaceous. Taking this live association and the depth limit of algal growth as food for the limpets into consideration, the mature ammonites dwelled or periodically visited the upper layer of the euphotic zone, probably less than around 20 m in depth. *** Ammonite, limpet, home depression, life mode, pseudoplankton, Cretaceous.     

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.     

Evidence for seasonal variation in δ13C and δ18O profiles of Baculites and implications for growth rate

The life histories of ammonites and the life strategies they employed are difficult to assess without robust modern analogues but placing constraints on ammonite growth rates provides a fundamental first step to understanding this abundant, but poorly understood, fossil group. Here we interpret periodic variations in carbon and oxygen stable isotope profiles from Campanian and Maastrichtian ammonites (Baculites) as seasonally driven and use these records to determine their rate of shell precipitation. Several of these samples are housed in museums and were originally prepared using sealants for display and preservation but testing of these sealants indicated no alteration of the isotopic values of treated carbonate. Diagenetic alteration, as determined by shell microstructure, affected the preservation of isotopic signals, resulting in the loss of seasonal variation in less well‐preserved specimens, and the δ¹³C signal is more robust than δ¹⁸O. The periodicity of isotopic profiles from Baculites shells presented here suggest that these organisms grew at rapid rates (c. 340 mm per year), which may imply an r‐type life strategy in which the animals reach maturity quickly, spawn large quantities of progeny, and die at a young age. Because of the potential mobility of Baculites, reconstructing palaeoenvironmental conditions from these isotopic records is challenging and should be conducted cautiously. Unfortunately, well‐preserved Baculites shells much longer than 350 mm are rarely recovered, which complicates the statistical treatment of potential periodicity in isotopic profiles.     

A new mode of ammonite preservation – implications for dating of condensed phosphorite deposits

Unusual phosphatic casts of the ammonites Mortoniceras (Subschloenbachia) sp. and Stoliczkaia sp. from the upper Albian condensed phosphorite bed at Annopol, Poland, are discussed in terms of their taphonomic history. These specimens are interpreted as ‘secondary’ external casts of ammonite replicas preserved originally as attachment scars on oyster shells. The following genetic history is suggested for this previously undocumented mode of ammonite preservation: (1) settling of shells of dead ammonites on the seafloor; (2) colonization of these shells by oysters and formation of ammonite replicas on left valves of oysters; (3) dissolution of ammonite shells; (4) reworking and fragmentation of oyster shells; (5) casting of ammonite replicas by phosphatic material; and (6) separation of ammonite casts from oyster shells, either through mechanical disintegration or dissolution of the latter. The specimens studied were formed after dissolution of the ammonite conchs, not prior to this event as in the case of typical ammonite steinkerns (internal moulds). Therefore, they are here referred to as ‘pseudo‐steinkerns’. The time interval between loss of the original ammonite shells and the formation of oyster‐mediated pseudo‐steinkerns may be very extensive. Therefore, the pseudo‐steinkerns may potentially mislead in biostratigraphic dating of condensed phosphorite deposits.     

An ammonite–fish association from the Kimmeridgian (Upper Jurassic) of La Rochelle, western France

A partially disarticulated actinopterygian fish preserved in a large three-dimensional ammonite body chamber is described from the Kimmeridgian of western France. Taphonomic observations on the degree of preservation of the fish and the development of epibiont organisms on the inner wall of the shell indicate a rather long time interval before sediment totally filled the body chamber. The fish, referred to an indeterminate Macrosemiidae, probably used this empty ammonite ( Rasenioides , Aulacostephanidae) shell as a refuge, or possibly for spawning and/or brooding. It can be assumed that ammonite shells may have constituted common shelters for demersal fishes living in an open-marine shelf environment, near to a muddy bottom devoid of rocks.     

Intraspecific variation of hatchling size in Late Cretaceous ammonoids from Hokkaido,Japan: implication for planktic duration at early ontogenetic stage

Tajika, A. & Wani, R. 2011: Intraspecific variation of hatchling size in Late Cretaceous ammonoids from Hokkaido, Japan: implication for planktic duration at early ontogenetic stage. Lethaia, Vol. 44, pp. 287–298. Intraspecific variations of the early shell dimensions (ammonitella and protoconch diameters) of two Late Cretaceous (earliest Campanian) ammonoid species (Gaudryceras tenuiliratum and Hypophylloceras subramosum) from the Haboro and Ikushumbetsu areas, Hokkaido, Japan, show no significant difference between these areas that are approximately 110 km apart. The geographic distributions of G. tenuiliratum and H. subramosum are supposed to be mainly controlled by the flotation and transportation during the embryonic stage within floating egg masses and/or post‐embryonic stage because of their small hatchling sizes (1.18–1.46 mm in diameter for G. tenuiliratum, and 0.91–1.13 mm in diameter for H. subramosum), suggesting these two ammonite species at the embryonic and/or post‐embryonic stages were transported at least 110 km. Postulating that the velocity of palaeocurrent around the Haboro and Ikushumbetsu areas during the Cretaceous Period was 0.25 m/s, similar to those in the modern ocean current flowing off the eastern Pacific coast of Hokkaido, the egg masses and/or hatchlings of G. tenuiliratum and H. subramosum were buoyant and transported more than 5 days. The preliminary comparison of hatchling size through time suggests that the hatching sizes of H. subramosum in Hokkaido increased slightly from the Middle Turonian until the earliest Campanian (during about 7 Myr). □ammonoid, hatchling, paleoecology, variation, Cretaceous.     

The oldest in situ hermit crab from the Lower Cretaceous of Speeton, UK

An Early Cretaceous (Hauterivian) hermit crab perfectly preserved within an ammonite is described. Palaeontologists have focused too much on the wrong class of molluscs in their attempt to analyse the evolution of the pagurids. The shift of mollusc habitation during the Cretaceous, from ammonites to gastropods, affected the morphology of the aperture–blocking hermit–crab claws.     

Mode of life and soft body shape of heteromorph ammonites

Ebel, K. 1992 04 15: Mode of life and soft body shape of heteromorph ammonites. Lethaia , Vol. 25, pp. 179–193. Oslo. ISSN 0024–1164.
Using the idea of a benthic mode of life for ammonites, based on a gastropod-like shell position, it is possible to reconstruct the development of all heteromorph ammonites by regarding single growth stages and the presumable acting forces. The reconstruction of shell formation, particularly the final shell position of heteromorphs with a hook, indicates that the soft body of the ammonite animal was considerably larger than comparison with the present-day Nautilus would suggest.     

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.     

First direct evidence of ammonoid ovoviviparity

Ammonoids had high evolutionary rates and diversity throughout their entire history and played an important role in the high‐resolution sub‐division of the Mesozoic, but much of their palaeobiology remains unclear, including the brooding habitat. We present our study of the first recorded ammonite embryonic shell clusters preserved with calcified embryonic aptychi in situ within the body chambers of mature macroconch shells of the Early Aptian (Early Cretaceous) ammonite Sinzovia sazonovae. The following support the idea that the clusters are egg masses, which developed inside ammonite body chambers: the absence of post‐embryonic shells and any other fossils in these clusters, the presence of the aptychi in all embryonic shell apertures and peculiarities of adult shells preservation. These facts confirm earlier speculations that at least some ammonoids could have been ovoviviparous and that, like many modern cephalopods, they could have reproduced in mass spawning events. The aptychi of ammonite embryonic shells are observed here for the first time, indicating that they were already formed and calcified before hatching. Our results are fully congruent with the peculiar modes of ammonoid evolution: quick recovery after extinctions, distinct evolutionary rates, pronounced sexual dimorphism and the nearly constant size of embryonic shells through ammonoid history. We assume that adaptation to ovoviviparity may be the reason for the presence of these features in all post‐Middle Devonian ammonoids.     

The jaw apparatus of the Cretaceous ammonite Reesidites

Jaws are preserved within the body chambers of three specimens of a collignoniceratid ammonite Reesidites minimus (Hayasaka and Fukada) from the Upper liuoniaq of Hokkaido, Japan. Light microscopic and SEM observations of sections indicate that both upper and lower jaws consist mainly of a thick, double-walled chitinous lamella with a beak-like anterior projection. The outer chitinous lamella of the lower jaw is covered by a thick calcareous layer. The jaw apparatus of this species morphologically resembles aptychus-type jaws of Jurassic ammonites, but is distinguished by the presence of an anterior beak-like projection with serrated ridges and grooves in the lower jaw. These observations strongly suggest a biting ability in this species.     

Ammoniten des Cenomans vom Südrand der westfälischen Kreide zwischen Unna und Möhnesee

Systematically collected ammonites from the different levels of the transgressive Cenomanian of the Haarstrang Mountains between Unna and Möhnesee (southern border of the westfalian Cretaceous, NW Germany) have been investigated. Twenty-one taxa (genera and species), partly previously unknown from this region, have been found and described. The fauna from the immediate transgression level is preserved as phosphoritic internal moulds (steinkerns). Fragmentary preservation and fossil condensation levels indicate reworking processes in these basal Cretaceous deposits. The age of the Cretaceous transgression in the area investigated can be precisely established as Lower Cenomanian (=carcitanensis subzone). The Cenomanian sequence, which is lithofacially very variable, can be, on the basis of the ammonite fauna, subdivided according to the modern Cretaceous stratigraphy for the boreal realm.     

Shell features, main colonized environments, and fractal analysis of sutures in Late Jurassic ammonites

A precise approach to the quantification of relationships between suture complexity, as measured by fractal analysis (step-line procedure), the architecture of shells, and the main colonized environments, has been made in a set of Late Jurassic ammonites ( N =507). Statistically significant differences between fractal-dimension ( D _f) mean values of evolute and involute shells are interpreted as caused by differences in the surface:volume ( S:V ) ratio. Suture complexity is also related to the shape of whorl section. The lowest D _f values correspond to subcircular whorl sections (low S:V ratio) and the highest ones to acute sections (high S:V ratio). The shape of flanks shows correlation with suture complexity. The highest values of D _f are found in planulate shells and the lowest ones in whorl cross sections with convex flanks. Highly significant differences appear between D _f mean values from unsculptured shells and those from ammonites with ribs and/or tubercles of medium to large size. Multivariate analysis shows a combined variation of shell features and suture complexity, resulting in a heterogeneous distribution of D _f values within the ammonite morphospace, mainly according to structural (shell architecture) and ornamental (sculpture strength rather than density) factors. Finally, the data obtained on relationships between suture complexity and the colonized environments (epicontinental vs. epioceanic inhabitants) suggest that suture complexity is not primarily related to bathymetry, and/or that no major differences in habitat depths existed between epicontinental and epioceanic ammonites.     

Commensal anomiid bivalves on Late Cretaceous heteromorph ammonites from south‐west Japan

The heteromorph ammonite Pravitoceras sigmoidale from the Upper Cretaceous Seidan Formation (Izumi Group) in south‐west Japan is frequently encrusted by sessile anomiid bivalves. Fossils of P. sigmoidale with anomiids are often concentrated at the top of or just above turbidite sandstones. Projecting retroversal hooks and apertures of P. sigmoidale are usually intact, and some individuals are associated with jaw apparatuses near apertures. Anomiids are found on both sides and ventral peripheries of P. sigmoidale conchs, attached predominantly to body chambers. These modes of occurrence suggest that the encrustation by anomiids occurred not on post‐mortem floating or sunken carcasses but on live conchs and that these organisms were rapidly buried by turbidity current deposits shortly after death. Attachment to both flanks and ventral peripheries of the retroversal hooks may indicate that at least adult individuals of P. sigmoidale did not lie on the sea floor and did not drag their body chambers. It is suggested that fully mature individuals of this ammonite species lived for a long period of time after having formed the retroversal hook because a few generations of anomiids have colonized a single body chamber. Such colonization by anomiids is also observed on Didymoceras awajiense, which is considered to be the closely related ancestral species of P. sigmoidale. This anomiid–heteromorph ammonite commensal relationship might continue to persist in descendants during the course of evolution of these heteromorph ammonites.     

Ammonite and inoceramid radiations after the Santonian-Campanian bioevent in Sakhalin, Far East Russia

Study of several marine Santonian-Campanian successions from Sakhalin Island, Far East Russia, has revealed that evolution of the ammonites and inoceramid bivalves proceeded at different rates after the major faunal turnover at the (locally defined) Santonian-Campanian boundary. Sometimes changes in the inoceramid assemblages were more frequent and rapid than changes in the ammonite assemblages, and sometimes vice versa. Significant levels of inoceramid turnover and radiation events have been identified at the (locally defined) Santonian-Campanian and lower-upper Campanian boundaries. Changes in ammonite and inoceramid diversity, and in the proportions of endemic and cosmopolitan species, were investigated in the context of the local relative sea-level curve and inferred environmental changes. In Far East Russia, the main ammonite and inoceramid radiation after the local Santonian-Campanian faunal turnover occurred in the early Late Campanian Pachydiscus (P.) aff. egertoni ammonite Zone and the coeval Schmidticeramus schmidti inoceramid Zone. This condensed interval of ammonite and inoceramid maximum diversity provides a perfect stratigraphic marker that is recognizable in Sakhalin, North-East Russia and Japan. The succession of Santonian-Campanian assemblages identified in Sakhalin enabled the establishment of seven ammonite and six inoceramid zones, which correlate relatively well with those of North-East Russia and Japan. The problems of placing the Santonian-Campanian boundary in Sakhalin and in the adjacent Japanese island of Hokkaido are reviewed.     

Sublethal injuries in Middle Jurassic ammonite shells from Poland

Sublethal injuries, the effects of which are seen as regeneration patterns, are described from Late Bajocian and Bathonian (Middle Jurassic) ammonites from Poland (Polish Jura area) for the first time. The total number of ammonite shells bearing signs of sublethal injuries is small (only 11 specimens, which constitute ∼1.2% of all ammonites investigated), and this value is even smaller (∼0.3 to 0.8%) when analysing a large sample of a particular ammonite species. Specimens under consideration represent ten species, belonging to six genera and five families. All the healed injuries are represented by only one type, referred to as the ‘forma verticata’ of Hölder. This type of regeneration, very common in ammonite shells in general, is an effect of a puncture injury of the shell-secreting mantle-epithelia at the apertural margin. Although many different extrinsic (mechanical) factors may be responsible for such healed injuries, here it is most plausible they are an effect of either competitive or predatory activities. Other causes, like collision of the ammonite shells with the substrate in a high-energy environment, are excluded because the sea-bottom was soft and situated below the storm wave-base. From many potential predators inhabiting the Polish Basin during the Bajocian and Bathonian, the most likely to have caused these injuries are other ammonites, belemnites and nautiloids. Crabs, which are cited in the literature as a probable perpetrator of the ‘forma verticata’ injuries, appear unlikely here, as the ammonites under discussion were not purely benthic.     

Palaeoecological significance of Barremian ammonite assemblages and facies variations from Southwest Mexico

Analyses of ammonite shell forms of two Barremian stratigraphic sections from Southwest Mexico consist of two well-defined morphotypes: (1) Small uncoiled, mostly leptoceratoid ancyloconic shells of the families Ancyloceratidae and Hamulinidae, and (2) middle-sized involute to moderately evolute oxycone to discocone shells of the family Pulchelliidae. Index taxa allow the recognition of standard ammonite biozones for the Barremian, which permit the relative dating of different processes that occurred through the water column in the environment of deposition. The vertical distribution of ammonite morphotypes and facies suggests changes of the palaeoceanographic and sedimentological conditions that prevailed in the area during Barremian time. Petrologic data, analyses of the organic carbon and carbonate contents of the rocks support the idea that oxygen-deficient bottom waters existed within a shallow marine, tectonically active area with little carbonate deposition during the early early Barremian (upper part of the Taveraidiscus hugii Zone through the base of the Nicklesia pulchella Zone). These conditions in the basin caused a proliferation of middle-water depth ammonites of Morphotype 1 but prevented the abundance of nektobenthic forms of Morphotype 2. Oxic conditions on a more calcareous and open normal marine environment seem to have been reestablished progressively during a transgressive episode from late early-early late Barremian (upper part of the Nicklesia pulchella Zone through the Gerhardtia sartousiana Zone). This environmental setting supported more facies dependent nektobenthic ammonites of Morphotype 2 to flourish within the basin.     

Nautilid nurseries: hatchlings and juveniles of Eutrephoceras dekayi from the lower Maastrichtian (Upper Cretaceous) Pierre Shale of east‐central Montana

The nautilid Eutrephoceras dekayi (Morton 1834) is relatively abundant in the lower Maastrichtian (Upper Cretaceous) Pierre Shale of east‐central Montana. We analysed the morphology, size frequency distribution, and isotope composition of a large collection of 220 well‐preserved specimens including hatchlings, juveniles and adults. The newly hatched shell is approximately 14 mm in diameter with a body chamber one‐third whorl in angular length terminating in the nepionic constriction. Internally, the embryonic shell contains five septa. Juveniles are abundant and comprise two‐thirds of the sample whereas sub‐adults, defined by the incipient flattening of the venter, are rare. Adults comprise approximately one‐third of the sample and average 100 mm in diameter. The co‐occurrence of newly hatched shells, small juveniles and adults suggests that the eggs were laid in the same area in which the hatchlings developed. Based on the excellent preservation of the juveniles, we conclude that they did not float into the area after death, but lived in the region, implying that this area served as a nursery for young animals. The calculated temperatures of the embryonic shells are similar to those of the post‐embryonic shells and generally range from 16 to 18 °C. Upon hatching, the nautilids probably followed a demersal mode of life and lived in well‐oxygenated water ≤50 m deep. An examination of lethal injuries (puncture holes) suggests that all ontogenetic stages were equally vulnerable to predation. The proximity of the site to the Sheridan Delta suggests that the specimens were smothered by sudden pulses of sediment transported into the area by major storms.     

The occurrence and preservation of ammonites in the Blue Lias Formation (lower Jurassic) of Devon and Dorset, England and their palaeoecological, sedimentological and diagenetic significance

More than two thirds of beds in the lowest Jurassic, Blue Lias Formation lack ammonites, which are commonly preserved in irregular or planar-bedded, bioturbated limestones, very rarely in laminated limestones and almost never in laminated black shales. Ammonites are preserved in 3D in nodular and planar-bedded limestones and at any orientation to bedding. Co-occurrence with macrobenthos and absence from beds without benthos suggest that Blue Lias ammonites were nektobenthonic. Scour structures and imbrication of ammonites in the Best Bed imply presence of traction currents. Lack of epifauna on large cephalopod shells (and other fossils) implies rapid deposition in event beds. Blue Lias deposition was episodic, not slow and continuous as the fine grain size implies. Undistorted trace fossils, uncrushed ammonites and stable isotope values all suggest early cementation of limestone beds from pore waters of a similar composition to contemporary Jurassic sea water. A clear diagenetic trend exists, with limestones having least, and laminated black shales most, modified stable isotope values. Contrast between trace fossil fills and host sediment demonstrates that Blue Lias rhythms are primary, but limestone beds have been diagenetically cemented.