The mode of life in ammonoids

The following structural features clearly indicate that ammonoid shells were adapted to withstand considerably higher hydrostatic pressures thanNautilus shells: (1) the corrugated and marginally fluted septa gave the shell wall efficient support against implosion; (2) the secondary connecting rings could grow a great deal in thickness; and (3) the last formed chambers remained full of liquid which supported the last septum. On the basis of the following characters it is concluded that ammonoids were incapable of swimming efficiently by jet-propulsion: (1) the retractor muscles were weakly developed; (2) the life position was unstable and highly variable; and (3) in animals with a ventral apertural rostrum the hyponome was probably absent. Ammonoids are considered here as having been pelagic cephalopods which lived in the upper 1000 m of the oceans, and which probably undertook considerable diurnal vertical migrations, similar to those inSpirula. Only some groups may have adopted a life in shallow epicontinental seas. In the late Mesozoic, ammonoids have been replaced by modern oceanic squids which are extremely numerous in the corresponding pelagic environment.     

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.     

On Frasnian ammonoids of the New Siberian Islands

Ammonoids from the basal beds of the Nerpalakh Formation (Lower Frasnian) of Belkovsky Island (New Siberian Archipelago) are systematically studied. Taxonomically, the assemblage studied (Manticoceras insulare sp. nov., Tornoceras typum (Sandberger, 1851), and T. contractum Glenister, 1958) is similar to the Early Frasnian ammonoid assemblage of South Timan, from which its is distinguished by the absence of the genera Timanites and Komioceras. The same beds contain conodonts of the Palmatolepis transitans Zone (= MN 4 Zone of the Montagne Noire standard succession), which allow the correlation of the beds studied with the Timanites keyserlingi and Komioceras stuckenbergi ammonoid zones of South Timan. The Early Frasnian ammonoids could supposedly have entered the region of the New Siberian Archipelago from the southwest at the time of a major transgression, which facilitated the distribution of the genera Manticoceras and Tornoceras. A new species of the genus Manticoceras is described.     

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.     

Ecological structure of Paleozoic ammonoids

Ecological specialization of Paleozoic ammonoid orders is discussed along with changes in the ecological structure of the ammonoid assemblages in the Early Devonian-Late Permian. Two large cycles of change in the ecological structure, Devonian and Carboniferous-Permian are recognized.     

Quantification of ontogenetic allometry in ammonoids

Ammonoids are well‐known objects used for studies on ontogeny and phylogeny, but a quantification of ontogenetic change has not yet been carried out. Their planispirally coiled conchs allow for a study of “longitudinal” ontogenetic data, that is data of ontogenetic trajectories that can be obtained from a single specimen. Therefore, they provide a good model for ontogenetic studies of geometry in other shelled organisms. Using modifications of three cardinal conch dimensions, computer simulations can model artificial conchs. The trajectories of ontogenetic allometry of these simulations can be analyzed in great detail in a theoretical morphospace. A method for the classification of conch ontogeny and quantification of the degree of allometry is proposed. Using high‐precision cross‐sections, the allometric conch growth of real ammonoids can be documented and compared. The members of the Ammonoidea show a wide variety of allometric growth, ranging from near isometry to monophasic, biphasic, or polyphasic allometry. Selected examples of Palaeozoic and Mesozoic ammonoids are shown with respect to their degree of change during ontogeny of the conch.     

Biogeographic evolution of Permian ammonoids

The distribution of Permian ammonoids within the five major biogeographic provinces (Arctic, Uralian, American, Tethyan, and Australian) is discussed. Changes in the taxonomic and biogeographic structure of assemblages according to stages in the evolution of Permian ammonoids are considered for each region. It is shown that the geographical distributions and taxonomic diversity of ammonoids changed continuously throughout the Late Permian.     

Permian ammonoids of the Kolyma-Omolon Region: Ogonerian association

The Ogonerian association of Early Permian ammonoids from the Upper part of the Munugudzhakian Formation (upper part of the Ogonerian Horizon) contains Uraloceras omolonense Bogoslovskaya et Boiko, U. kolymense Bogoslovskaya et Boiko, Neoshumardites munugudzhensis sp. nov., N.? nassichuki sp. nov., and Bulunites gracilis sp. nov. These taxa suggest the Sakmarian-Artinskian boundary beds examined are geochronologically older than the level with Neoshumardites triceps Ruzhencev in the South Ural Mountains.     

Soft-part anatomy of the siphuncle in Permian prolecanitid ammonoids

Exceptionally well-preserved remains of phosphatized siphuncles were discovered in four specimens of a Permian prolecanitid ammonoid Akmilleria electraensis (Plummer and Scott) from Buck Mountain, Nevada. These structures occur as truncated segments within the siphuncular tube. The outer surface of the siphuncle is sculptured by numerous equally spaced longitudinal ridges and furrows; the ridges represent an infolded basement membrane of epithelial cells which corresponds to the distal ends of individual canaliculi between epithelial cells. In cross-section, the siphuncle of A. electraensis consists of a large central vein, possibly two pairs of arteries, porous connective tissue with reticulate hemocoelic spaces, and a thin epithelium. In the presence of two pairs of arteries and porous connective tissue, the siphuncle of A. electraensis is more like that of Nautilus pompilius than that of Spirula spirula, which has nine arteries and dense connective tissue. However, Nautilus possesses relatively smaller and more numerous epithelial cells around the siphuncle than does Akmilleria. These observations strongly suggest that the siphuncular epithelium of Akmilleria served as the salt-­concentrating organ for buoyancy regulation of the living animal, just as in Nautilus and Spirula.     

The A-mode sutural ontogeny in prolecanitid ammonoids

The generally accepted theory of a U-mode sutural ontogeny of prolecanitid ammonoids and their descendants is refuted. The basic suture formula of the order Prolecanitida is E A L U I (not E L U₂ U₁ I), resembling that of derived members of the suborder Tornoceratina (their phylogenetic ancestors), and of the suborder Goniatitina. During phylogeny of the prolecanitids, secondary umbilical lobes are introduced and lead to multilobate forms. In early ceratites, the original L lobe disappeared, and an increase in sutural elements took place by the introduction of supplementary U lobes. Consequently, sutural nomenclature of Permian ceratites and Mesozoic ammonoids has to be modified.     

Role of heterochronies in the morphogenesis of paleozoic ammonoids

Various types of evolutionary changes related to shifts in the appearance or order of formation of characters, i.e., heterochronies are shown. The most common such changes in Paleozoic ammonoids was the appearance of mosaic forms combining conservative and advanced characters resulting from recapitulation and paedomorphosis. These phenomena are related to certain phylogenetic stages in each group: the growth stage predominantly showed recapitulations; the diversification stage showed the appearance of many mosaic forms; the decline stage commonly shows paedomorphosis connected with retardation.     

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.     

Permian ammonoids of the Kolyma-Omolon Region: Kyrian association

This paper describes new taxa of the earliest Permian ammonoids of the Kolyma-Omolon Region: the genus Kolymoglaphyrites gen. nov. with the type species K. lazarevi sp. nov. and the species Uraloceras margaritae sp. nov., recognized as the Kyrian ammonoid association. Judging from the analysis of the main characters, the genus Kolymoglaphyrites is included in the mainly Carboniferous family Glaphyritidae, which expands the interval of the vertical distribution of this family to include the Sakmarian. Uraloceras margaritae is considered as the initial genus in the lineage of weakly ornamented species of Uraloceras. Apparently, this species evolved from Paragastrioceras sterlitamakense at the Asselian-Sakmarian boundary to become later ancestral for U. simense. Based on the phylogenetic position of U. margaritae the Kyrian ammonoid association is dated as Early Sakmarian.     

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.     

The genus Luppovia and the phylogeny of Cretaceous heteromorphic ammonoids

The unique preservation of heteromorphic shells of Luppovia Kakabadze, Bogdanova & Mikhailova 1978 from the Aptian deposits of Bolshoj Balkhan (western Turkmenia) made it possible to study the microstructure, internal shell characteristics and suture ontogeny of this genus. Microstructural investigations revealed that the dorsal shell wall in Luppovia is composed of three layers in contrast to that of monomorphic ammonoids, which is single-layered. This ability of secreting different numbers of layers in monomorphs and heteromorphs demonstrates plasticity of secreting activity of the mantle epithelium. Data obtained from the study of internal shell structure and suture ontogeny have been compared. Both these methods of investigation lead to the conclusion that the genus Luppovia belongs to the order Ammonitida. The results obtained confirm the viewpoint that the Cretaceous heteromorphic superfamilies Turrilitaceae and Ancylocerataceae have independent origins and belong to two different orders, Lytoceratida and Ammonitida, respectively. □ Ammonoidea , Luppovia, shell structure, ontogeny, phylogeny, Cretaceous.     

The connecting rings of Nautilus and Mesozoic ammonoids: implications for ammonoid bathymetry

The fresh conchiolinous ('horny') connecting rings of Nautilus pompilius decrease ontogcnctically in the strength index (100 times; wall thicknessh/inner radius r), i.e. from ≥ 16 to – 12. A similar decrease of 100 h/ r is the rule in Ammonitina. This may reflect depth migration but, more likely and in analogy with Nautilus, is compensated for by increased conchiolin strength. The mature ammonoid siphunclc, therefore, provides the more reliable indicator for depth limits. No marked shrinkage of connecting rings is evident in fossil nautiloids, but Mesozoic nautilid connecting rings are rarely preserved. Post-mortem alteration of the abundantly preserved ammonoid connecting rings entails mechanical distortion and fragmentation by differential filling of the siphunclc and camerac with sediment, and probable microbial decomposition, proceeding from the body chamber and resulting in pitting and perforation; shrinkage occurred rarely. This is probably related to the high phosphate content as here documented for Haplo-phylloceras. If the phosphate prevalent in ammonoid connecting rings is a primary constituent, rather than of early diagenetic origin, calibration of the ammonoid strength index on the non-phosphatic Nautilus connecting rings may not be justified. The strength indices of ammonoid connecting rings would thus provide only a relative scale of depth limits. Another example of a phylloccratid, Ptychophylloceras, has been found with a connecting ring extending well into the body chamber. D Connecting rings, ammonoids. Nautilus, bathymetry.     

Major trends in the evolution of Permian ammonoids

The phylogeny of major families of Permian ammonoids is analyzed. The evolution of most families followed a typical scenario with distinct stages of early evolution, diversification, and decline. A smaller group followed a different evolutionary narrative, with indistinct stages. The former group includes families with both simple and complex morphology and a wide range of variation. The nature and trends in the evolution of the families may change depending on their phylogenetic stage. The Early Permian (Asselian), the second half of the Artinskian, and the beginning of the Middle Permian were marked by the most significant evolutionary changes. The Late Permian was the time of the decline of Paleozoic ammonoid orders and of the onset of the evolution of the Mesozoic order Ceratitida.     

New middle Olenekian (early Triassic) ammonoids of South Primorye

Nine new ammonoid species (Inyoceras singularis, Yvesgalleticeras proximus, Tirolites opiparus, Koninckitoides solus, Bajarunia magna, Albanites vulgaris, Nordophiceratoides praecox, Palaeophyllites admirandus, Kamenushkaites acutus) and one new genus of the family Palaeophyllitidae (Kamenushkaites) are described based on material from the mid-Olenekian of the Kamenushka River basin, South Primorye.     

Key innovations in Mesozoic ammonoids: the multicuspidate radula and the calcified aptychus

A nearly complete radula with seven elements per row preserved inside of an isolated, bivalved, calcitic lower jaw (= aptychus) of the Late Jurassic ammonite Aspidoceras is described from the Fossillagerstätte Painten (Bavaria, southern Germany). It is the largest known ammonite radula and the first record for the Perisphinctoidea. The multicuspidate tooth elements (ctenodont type of radula) present short cusps. Owing to significant morphological differences between known aptychophoran ammonoid radulae, their possible function is discussed, partly in comparison with modern cephalopod and gastropod radulae. Analogies between the evolution of the pharyngeal jaws of cichlid fishes and the ammonoid buccal apparatus raise the possibility that the evolution of a multicuspidate radula allowed for a functional decoupling of the aptychophoran ammonoid jaw. The radula, therefore, represents a key innovation which allowed for the evolution of the calcified lower jaws in Jurassic and Cretaceous aptychophoran ammonites. Possible triggers for this morphological change during the early Toarcian are discussed. Finally, we hypothesize potential adaptations of ammonoids to different feeding niches based on radular tooth morphologies.