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In morphological studies the shape may be conveniently quantified by relative dimensions or dimensionless quantities. The analyses of shell morphology and morphospace occupation have been historically approached mainly by means of statistical analysis on classical dimensions (distance measurements: diameter, umbilical width, whorl width, whorl height and apertural whorl height), the Raup’s coiling and expansion rate parameters and, more recently, by means of the ADA−model which conjugates the classical variables in a single simple equation. Relationships between these studies should be possible based on mathematical equivalences between classical dimensions and those of coiling and expansion rates. These equivalences, which are presented in this paper, have been obtained on the basis of the ADA−model and a new general method for deriving dimensionless models of morphology based on exponential trajectories as a function of a rotational angle.
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The study of pristine preserved shells of Mesozoic Ammonoidea shows different types of construction and formation of the dorsal shell wall. We observe three major types: (i) The vast majority of Ammonoidea, usually planispirally coiled, has a prismatic reduced dorsal shell wall which consists of an outer organic component (e.g., wrinkle layer), which is the first layer to be formed, and the subsequently formed dorsal inner prismatic layer. The dorsal mantle tissue suppresses the formation of the outer prismatic layer and nacreous layer. With the exception of the outer organic component, secretion of a shell wall is omitted at the aperture. A prismatic reduced dorsal shell wall is always secreted immediately after the hatching during early teleoconch formation. Due to its broad distribution in (planispiral) Ammonoidea, the prismatic reduced dorsal shell wall is probably the general state. (ii) Some planispirally coiled Ammonoidea have a nacreous reduced dorsal shell wall which consists of three mineralized layers: two prismatic layers (primary and secondary dorsal inner prismatic layer) and an enclosed nacreous layer (secondary dorsal nacreous layer). The dorsal shell wall is omitted at the aperture and was secreted in the rear living chamber. Its layers are a continuation of an umbilical shell doubling (reinforcement by additional shell layers) that extends towards the ventral crest of the preceding whorl. The nacreous reduced dorsal shell wall is formed in the process of ontogeny following a prismatic reduced dorsal shell wall. (iii) Heteromorph and some planispirally coiled taxa secrete a complete dorsal shell wall which forms a continuation of the ventral and lateral shell layers. It is formed during ontogeny following a prismatic reduced dorsal shell wall or a priori. The construction is identical with the ventral and lateral shell wall, including a dorsal nacreous layer. The wide distribution of the ability to form dorsal nacre indicates that it is a plesiomorphic trait which either was passed on from gyrocone ammonoid ancestors or (re-)developed in post-Triassic ammonoids.
The most distinctive and important element of the hydrostatic organ of ammonoids and nautiloids is the siphuncular tube. It consists of mineral and organic segments (so−called connecting rings). The connecting ring of ammonites never preserves its original organic matter in the mineralized state, usually having undergone diagenetic phosphatisation, more rarely, calcification, or even complete loss. Our knowledge about its original ultrastructure is based upon comparison with Recent Nautilus and phosphatised or calcified ammonite fossils. We show that depending on the taphonomic history, both calcium phosphate and calcite can participate in the diagenesis of the connecting ring wall. Under standard light microscopy, the phosphatised elements are indistinguishable from the calcified ones. Both are dark brown in colour, due to an excess of carbon. The structure of the phosphatised siphuncle does not closely replicate the structure of its organic elements. This casts doubts on conclusions of other authors who described a complex porous structure in ammonite siphuncles, which is completely dissimilar to the siphuncular structure of Recent Nautilus and suggests that this organ functioned differently in ammonites. SEM observations using a BSE detector on the calcified parts of the walls of connecting rings revealed a multilayered structure with perpendicular elements connecting particular layers, resembling the structure of a stacked nacreous layer.
Ammonoid biodiversity changes from shallow to offshore environments across the Cenomanian–Turonian (C–T) boundary are reconstructed in the Yezo Group, Hokkaido, Japan. This group was probably deposited at approximately 35–45ºN along a westward subduction margin in the northeastern Asian continent. Temporal changes in species richness in the Yezo Group, which show persistently high values during the middle Cenomanian and then decline stepwise from near the middle–late Cenomanian boundary, resemble those in Europe, but not those in Tunisia and the Western Interior. These differences suggest that the Cenomanian–Turonian “mass extinction” was not a global event for ammonoids but was restricted to mid−palaeolatitudinal regions (Europe and Japan). Sea level and climate changes probably influenced ammonoid faunas in the Yezo Group as well as those in Europe. However, it is unlikely that a single, simple cause led to the C–T boundary “mass extinction” because various abiotic changes in the Cenomanian–Turonian transition have been detected, and biotic and abiotic change are interrelated.
Early late Viséan ammonoid assemblages in Morocco are composed of diverse and well-preserved specimens. The material was found in a plain in the Tafilalt (eastern Anti-Atlas). Here, we describe mass-occurrences of juvenile specimens, in which subadult and adult specimens occur in low numbers. The juveniles of some species display a conch morphology that differs fundamentally from the adult stages. Accordingly, we emend the species diagnoses of Goniatites lazarus as well as Calygirtyoceras darkaouaense, introduce the species Entogonites bucheri sp. nov., and discuss possible ecological implications of the morphologic changes throughout ontogeny. In particular, we compare the changes in conch morphology through ontogeny in the light of Pareto Optimiality according to which the morphology of organisms would fill a polygon or polyhedron in morphospace. Data points in one of the vorteces of the polyhedron indicate optimisation for the corresponding task. Although shape is not a proof of function, it appears plausible that juvenile conchs were selected rather for compactness while adult conchs were positively selected for conchs with improved hydrodynamic properties. This appears plausible because at small conch diameters, swimming movements will not suffice for effective translocation and a planktonic mode of life is likely.
The Middle Triassic ammonoid genus Ceratites diversified spectacularly within the Germanic Muschelkalk Basin during the Anisian/Ladian (244–232 Mya). Previous studies have interpreted this diversification as a sequence of rapid, endemic radiations from a few immigrant taxa. Here we investigate the possibility that geological and sampling biases, rather than ecological and evolutionary processes, are responsible for this pattern. A new specimen−based dataset of Ceratites species−richness and abundance was assembled. This dataset was combined with 1:200 000 geological maps in a geodatabase to facilitate geospatial analyses. One set of analyses compared species richness per geological map with the number of occurrences and localities per map. Per−map change in the amount of rock available to sample for fossils was also included as a variable. Of these three variables, number of occurrences is the most strongly correlated with richness. Variation in the amount of rock is not a strong determinant of species−richness. However, rarefaction of basin−wide species/abundance data demonstrates that differences in species−richness through time are not attributable to sample size differences. The average percent similarity among sites remained close to 50% throughout the Upper Muschelkalk. The rank abundance distribution (RAD) of species from the first interval of the Upper Muschelkalk is consistent with colonization of a disturbed environment, while the other two intervals have RADs consistent with more stable ecosystems. These results indicate that genuine ecological and evolutionary events are partly responsible for the observed differences in richness and abundance. Although changes in the RADs through time support changes in the ammonoid assemblage structure, the processes underlying increasing richness and change in RADs cannot be explained by increasing geographic distinctiveness or isolation among the ammonoid assemblages present at different localities.
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The origin of ammonoid locomotion

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Evolution of the coiled ammonoid conch from the uncoiled bactritid conch was probably coupled with changes in manoeuvrability and swimming velocity. The gradual transformation of uncoiled to coiled ammonoid conchs has essential functional consequences. The radical change in conch geometry during phylogeny but also in ontogeny of early ammonoids implies a shift of the aperture from an original roughly downward, via a downward oblique and an upward oblique to an upward orientation, presuming a neutrally buoyant condition of the ammonoid animal. Similar trends were reconstructed for the three main ammonoid lineages in the Middle Devonian, the agoniatitid, the anarcestid, and the tornoceratid lineages. This allowed an increase in manoeuvrability and in the maximum horizontal swimming speed.
Predatory shell breakage is known to occur occasionally on the ventrolateral portion of the body chamber in Mesozoic ammonoids. Here we report, for the first time, quantitative data of shell breakage in large ammonoid samples that were recovered from the lower Toarcian (Lower Jurassic) strata in the Toyora area, western Japan. The strata yielding the ammonoid samples consisted mostly of well-laminated, bituminous black shale that was deposited in an oxygen-depleted shelf basin of the northwestern Panthalassa, under the influence of the early Toarcian oceanic anoxic event. Among a total of 1305 specimens from 18 localities, apparent shell breakage was recognised in 35 specimens belonging to 7 genera, resulting in only a 2.7% frequency of occurrence relative to the total number of specimens. The breakage occurs mostly on the ventrolateral side of the body chamber with a complete shell aperture. This fact, as well as the low energy bottom condition suggested for the ammonoid-bearing shale, indicate that the shell breaks observed in the examined ammonoids were not produced by non-biological, post-mortem biostratinomical processes but were lethal injuries inflicted by nek-tonic predators such as reptiles, jawed fishes, coleoids, nautiloids, and carnivorous ammonoids with calcified rostral tips in their upper and lower jaws. Similar predatory shell breaks on the ventrolateral side of the body chamber have been found in contemporaneous ammonoid assemblages of the Tethys Realm, with a much higher frequency of occurrence than in the examined samples from the northwestern Panthalassa, suggesting a weaker durophagous predation pressure on ammonoids in the latter bioprovince.
Investigations of the Upper Permian strata in the Iran−Transcaucasia resulted in identification of 32 ammonoid genera. The majority of ammonoids in this collection belong to the order Ceratitida (75%). Among Dzhulfian ceratitid ammonoids representatives of the family Araxoceratidae (Otoceratoidea) are most abundant. The assemblage structure changed radically during latest Permian (Dorashamian) time, bringing a domination of the family Dzhulfitidae. The Induan (Lower Triassic) succession in the Verkhoyansk area provided a few groups of ammonoids which are Palaeozoic in type: families Episageceratidae (Episageceras), Xenodiscidae (Aldanoceras and Metophiceras), and Dzhulfitidae (Tompophiceras) and superfamily Otoceratoidea (Otoceras and Vavilovites). It demonstrates the survival of ammonoids belonging to these groups the Permian–Triassic (P–T) boundary extinction event and their quick migration to the vast ar− eas of higher latitudes (together with some representatives of the Mesozoic−type families). Induan–Olenekian ammonoid successions in South Primorye, Mangyshlak, and Arctic Siberia illustrate the high rate of Early Triassic ammonoid recov− ery in both the Tethys and the Boreal realm. New ammonoid taxa are described: Proptychitina subordo nov., Ussuritina subordo nov., Subbalhaeceras shigetai gen. and sp. nov. (Flemingitidae), Mesohedenstroemia olgae sp. nov. (Heden− strormiidae), and Inyoites sedini sp. nov. (Inyoitidae).
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Raised shell projections on the inner shell walls that form pits on the internal moulds of Devonian ammonoids have been known for several decades. New specimens from Morocco reveal novel details of these structures; most, if not all, of which consist of a capsule of ammonoid shell that covers tiny tubes attached to the outer (= lateral or ventral) shell wall from the inside. In accordance with comparable Recent occurrences of similar structures in molluscs, we use the term “pearls” for these structures and the pits they form on the internal moulds. The nature of these encapsulated tubes is described and discussed. Because of the presence of these tubes inside the pearls, pearl arrangement, and their similarity to Recent mollusc occurrences, the tubes are interpreted as traces of parasitoses. The pearls and pits were grouped into five types based on differences in morphology, size, and arrangement. Then, having used these traits to perform a simple cladistic analysis, the resulting cladogram was compared to the phylogeny of ammonoids. Based on this comparison, it appears likely that the parasites underwent a co−evolution with the ammonoids, which lasted 10 to 15 Ma. Patterns of evolutionary events include co−speciation, “drowning on arrival” (end of parasite lineage near base of a new host clade), and “missing the boat” (parasite lineage does not adapt to a new host clade, thus not evolving a new parasite clade). Because of the lack of fossilised soft tissue, only speculations can be made about the systematic affiliation of the parasites, their life−cycle, infection strategy, and ecological framework. Some co−occurring bivalves also have pits reminiscent to structures caused by trematodes in Recent forms. Based on the available information, the tubes are interpreted as artefacts of trematode infestations, which, if correct, would extend the fossil record of parasitic trematodes into the Early Devonian.
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Dorsal shell wall in ammonoids

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In ammonoids, a soft body organ (possibly a supracephalicmantle fold), extending from the conch aperture secreted aragonitic wrinkles, forming a layer on the surface of the preceding whorl. The dorsal shell wall consists of the outer and inner components which were deposited sequentially, beginning at the aperture of the living chamber inwards. The dorsal wall attains its full thickness near the last septum. The outer component is visible in the apertural region and is smooth or wrinkled; it is called the wrinkled layer in the latter case. The wrinkles may be continuous, interrupted, or form isolated patches arranged in rows. The wrinkles are usually triangular in cross section. A further stage of dorsal wall development involves filling in the space between the apices of triangles, and then adding one or more inner prismatic layers from the inside of the living chamber. This pattern occurs at least in the postembryonic stage of all genera studied, belonging to five suborders of Ammonoidea ranging from Late Carboniferousto Late Cretaceous. In many genera, the outer component of the dorsal shell wall exhibits remarkable ontogenetic change in its ultrastructure and microornament. It may be compared with the black film of Recent Nautilus shells with respect to place of formation. The outer component of the ammonoid dorsal shell wall is regarded as a product of organic secretion and carbonate precipitation in the area of the supracephalic mantle fold.
Organic membranes preserved in the rear part of the body chamber of the Late Cretaceous phylloceratid ammonite Phyllopachyceras ezoense were examined with scanning electron microscopy (SEM) on the basis of well−preserved specimens from Hokkaido, Japan. SEM observations revealed that the membranes are continuous with the siphuncular tube wall in the phragmocone and consist of two layers, both of which are made of a dark, primarily conchiolin material; namely, a thinner inner homogeneous layer and a thicker outer layer with gently inclined pillar−like units. Hence, they are interpreted as the precursory siphuncular membranes. The precursory siphuncular membranes are not associated with any other organic components such as the siphuncular sheets reported in some Paleozoic and Mesozoic ammonoids. Unlike the tube−like condition in the phragmocone, the precursory siphuncular membranes in the body chamber of the specimens examined do not form a tube shape; on the ventral side the membranes are truncated and directly contact the outer shell wall. These observations suggest that the inner and outer layers of the precursory siphuncular membranes in the body chamber were respectively formed by the siphuncular epithelium from the inner side and by the invaginated septal epithelium from the outer side. It is also postulated that at the initial stage of septal formation, the rear part of the body moved slowly forward, developing a circumsiphonal invagination of the septal epithelium. Because similar conchiolin membranes are occasionally preserved in the body chambers of other phylloceratids, the above morphogenetic process applies to all members of the Phylloceratina. The tube−shaped structure in the rear part of the body chamber of desmoceratid Damesites consists only of nacreous layer. We interpret it as a pathologically overgrown prochoanitic septal neck.
The dominant factor in faunal succession of conodonts in the Frasnian of Poland is the apparent immigration of species originating allopatrically in other regions. Each immigration event usually changes the population variability of a local species (character displacement). Only a few lineages show their phyletic evolution within the studied area. Attempts to distinguish conodont species on the basis of platform element shape failed in some of the latest Frasnian palmatolepidids. Even at the apparatus−based generic level, certain ramiform elements of the apparatus appear much more diagnostic than the platforms. Correlative value of the late Frasnian palmatolepidids of unknown apparatus structure is thus questionable. The evolution of platform elements in Ancyrodella offers a more solid basis for age determination although their great population variability makes resolution rather low and requires the population approach. The panderodontids Belodella(?) tenuiserrata sp., B. minutidentata sp. nov., B. robustidentata sp. nov., prioniodontid Icriodus kielcensis sp. nov., enigmatic monospecific Playfordiidae fam. nov., prioniodinids Dyminodina planidentata gen. et sp. nov., D. anterodenticulata sp. nov., D. kovalensis sp. nov., Pluckidina kielcensis gen. et sp. nov., P. slupiensis sp. nov., P. robustipegmata sp. nov., and P. lagoviensis sp. nov., derived polygnathid Avignathus bifurcatus sp. nov., probably secondarily simplified polygnathid Nicollidina gen. nov., and palmatolepidids Kielcelepis gen. nov., Lagovilepis gen. nov. and Klapperilepis gen. nov. are proposed.
The Shemshak Group at Shahmirzad (northern Iran) is characterized by the most frequent and extensive marine intercalations and contains the most abundant and diverse ammonite faunas hitherto known from the Lower and lower Middle Jurassic strata of the Alborz Range. So far, 62 ammonite taxa have been recorded from this area, including 25 taxa from earlier studies. The taxa belong to the families Cymbitidae, Echioceratidae, Amaltheidae, Dactylioceratidae, Hildoceratidae, Graphoceratidae, Hammatoceratidae, Erycitidae, and Stephanoceratidae with the new species Paradumortieria elmii and Pleydellia (P.?) ruttneri. The fauna represents the Late Sinemurian, Late Pliensbachian, Toarcian, Aalenian, and Early Bajocian. Palaeobiogeographically, it is closely related to the Northwest European (Subboreal) Province, and exhibits only minor relations with the Mediterranean (Tethyan) Province.
A new preservation of aptychi is described from the Valanginian limestone−marl alternations of the Vergol section (Drôme), located in the Vocontian Basin (SE France). Aptychi are arranged into two parallel rows which are generally 50 mm in length and separated by 4 mm. The alignments are very often made by entire aptychi (around 10 mm in length), oriented following their harmonic margin. Aptychi show the outside of valve to the viewer: they are convex−up. This fossilization of aptychi is successively interpreted as the result of post−mortem transport by bottom currents (taphonomicresedimentation process) or the residues (bromalites: fossilized regurgitation, gastric and intestinal contents, excrement) from the digestive tract of an ammonoid−eater (biological processes). Both the parallel rows of aptychi are more likely interpreted as a coprolite (fossil faeces) and they could be considered as both halves (hemi−cylindrical in shape) of the same cylindrical coprolite which would have been separated in two parts (following the long axis) just after the animal defecated. Considering this hypothesis, a discussion is proposed on the hypothetical ammonoid−eater responsible for them.
This paper reports on the evolution of ammonoids belonging to the family Tornoceratidae from the Devonian of Janczyce in the Holy Cross Mountains, Poland. Steady and gradual changes in conch morphology of the goniatite lineage Phoenixites frechi–Tornoceras subacutum–T. sublentiforme occurred in concert with water shallowing during the deposition of the Lower Famennian cephalopod limestone. Biometric analysis of ammonoid conch and facies analysis of the cephalopod limestones have been applied to assess the possible relationship between shell geometry and environmental changes. Results show that ratios of whorl width / diameter as well as whorl width / whorl height decreased, while distance from the venter to the greatest whorl width / diameter increased with time, thereby reducing hydrodynamic drag of the shells, probably in response to increasing water turbulence. The interpretation presented here is in agreement with similar cases from the literature. However, this kind of environmentally controlled evolution has hitherto been recognized only in Jurassic and Cretaceous ammonoids. Conch morphology may be considered as an indicator of palaeobathymetry.
A new theoretical morphological model is proposed for the analysis of growth, form and morphospace of ammonoid shells. In this model, the shape of a radial cross section through the shell is simulated by “piggybacking” of successive whorls. The “piggyback whorls model” is defined in terms of the enlarging rate of the perimeter and the proportion of the dorsal wall to the whorl periphery, if an isometric relationship is assumed between perimeter and area of the cross−sectioned whorl. Allometric coefficients on these growth parameters determine how compressed and evolute shells are formed. The present model successfully reproduced some correlations among purely geometric variables that have been reported in previous works and were also observed in our biometric analyses. This model yields a hypothesis of “constructional linkages” between aperture shape and coiling geometry that might provide a functional coupling between hydrostatic and hydrodynamic characters. The model may partly explain Buckman’s Law of Covariation between rib features and shell shapes.
The taphonomy of the Upper Cretaceous ammonite Anagaudryceras limatum differs from associated ammonites in the same horizon. This differential taphonomy is reconstructed based on fragmentation patterns and structural property of the corrugated shells of their body chamber parts. The characteristic preservation of isolated body chamber parts with corrugations is commonly observed in A. limatum from the upper Turonian of the Oyubari area, central Hokkaido, Japan. This preservation probably results from the differential internal structural properties of the hollow body chamber and septated phragmocone as well as the peculiar corrugation on body chamber of the present species: (1) fracturing along the corrugations against bending force is easier than flat material, because of the concentration of force on the tops or bottoms of corrugations, and (2) high durability against compressive forces. The separated body chamber parts were resistant to being squashed and broke into pieces due to the durability of corrugations against compressive force. The statistic test on the fossil assemblage suggests that selective destruction did not affect the fidelity of species composition and relative abundance of the studied fossil assemblage. Molluscan death assemblages in marine systems consistently show strong fidelity to relative abundances in the live community, suggesting that there was an Anagaudryceras−dominant ammonoid community during the late Turonian in the Oyubari area.
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