Aleksandr Mironenko - Academia.edu (original) (raw)

Papers by Aleksandr Mironenko

Research paper thumbnail of The first record of jaws of Boreal Valanginian ammonites (Cephalopoda, Polyptychitidae

Jaws of ammonites which inhabited the Panboreal Superrealm during the Jurassic and Cretaceous are... more Jaws of ammonites which inhabited the Panboreal Superrealm during the Jurassic and Cretaceous are poorly known in comparison to those of Tethyan ammonoid faunas. This paucity may be explained by limited thickness, or even absence of an outer calcitic layer, in lower jaw elements (aptychi) of Boreal ammonites. Here we describe, for the first time, the jaws (both lower and upper) of ammonites of the Boreal family Polyptychitidae, of Early Cretaceous (Valanginian) age. Polyptychitid lower jaws are of the aptychus type, but have an unusual pointed and convex shape. However, lower jaws of Late Jurassic ancestors of polyptychitids (Craspeditidae) as well as Middle Jurassic cardioceratids (Pseudocadoceras) have a near-identical shape, as do previously described aptychi of the Late Cretaceous genera Neogastroplites and Placenticeras (Hoplitoidea). The close resemblance of lower jaws of evolutionarily distant ammonites may be linked to a similar lifestyle, but more data are needed to substantiate this. Upper jaws of polyptychitid are closely similar to previously described upper jaws of Jurassic ammonites, which indicates the conservatism of this part of the jaw apparatus. Together with shells and jaws of the Valanginian ammonites described herein, jaws of coleoids (likely belemnites) as well as arm hooks (onychites) have been found.

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Research paper thumbnail of First discovery of the soft‐body imprint of an Oligocene fossil squid indicates its piscivorous diet

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Research paper thumbnail of Structure of the calcitic layer of the aptychus of the ammonite genus Peltoceras

Annales Societatis Geologorum Poloniae, 2021

The aptychi of ammonites combined the functions of lower jaws and protective opercula. They consi... more The aptychi of ammonites combined the functions of lower jaws and protective opercula. They consist of two parts: an inner organic layer and an outer calcitic lamella. In different evolutionary lineages of ammonites, the shape of aptychi, the sculpture of their surface and the microstructure of the calcitic layer vary greatly. However, the structure of the aptychi is not known for all evolutionary lineages of ammonites. Although numerous aptychi have been described for the Jurassic family Aspidoceratidae, almost all of them belong to only one evolutionary branch of this family – the Aspidoceratinae (sensu lato). For the second branch – the Peltoceratinae, only one aptychus had been described to date and the structure of its calcitic layer remained unknown. In this article, for the first time, the structure of the aptychus of the Peltoceratinae (upper Callovian Peltoceras) is described. The surface of this aptychus is covered with rough ribs and the calcitic part consists of only one layer of dense calcite. The thickness of the aptychus is much greater than that of the aptychi of supposed ancestors of the Peltoceratinae. The increase in the thickness of the aptychi in both the Aspidoceratinae and the Peltoceratinae, contemporaneously with the appearance of spines on their shells, is most likely related to increasing the protective function of the aptychi of these ammonites in the late Callovian.

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Research paper thumbnail of An unusual conchorhynch from the upper Maastrichtian of the southeast Netherlands and the distinction between nautiloid and ammonoid conchorhynchs (Mollusca, Cephalopoda)⁋

Cretaceous Research, 2022

A single, atypical conchorhynch (calcitic tip of a cephalopod lower jaw), recovered from the uppe... more A single, atypical conchorhynch (calcitic tip of a cephalopod lower jaw), recovered from the uppermost Meerssen Member (Maastricht Formation, upper Maastrichtian) at the former ENCI-HeidelbergCement Group quarry, south of Maastricht, is described as a new parataxon, Conchorhynchus illustris sp. nov. The specimen can be differentiated from all previous conchorhynch records on account of its large size, elongated shape and, in particular, of the structure of its apical part which is smooth and forwardly elongated. During the Late Cretaceous, conchorhynchs formed part of the jaw apparatus of nautilids and of two ammonoid suborders, Phylloceratina and Lytoceratina. Since conchorhynchs are most often found separated from jaws, establishing to which group of cephalopods their bearer belonged can be complicated. Here, for the first time, we propose a set of morphological criteria to differentiate clearly between nautiloid and ammonoid conchorhynchs. Although Conchorhynchus illustris sp. nov. is distinct from all currently known nautilid conchorhynchs, the sum of its morphological features is indicative of assignment to that cephalopod group. The upper portion of the Maastricht Formation in the Maastricht area (Nekum and Meerssen members) has yielded internal and external moulds of shells of the nautilid Eutrephoceras and the hercoglossid Cimomia. The new conchorhynch type described herein most likely belonged to one of these shell-based taxa. Judging from its unusual shape, the feeding strategy of its bearer must have differed from that of modern nautilids, in that it held and pierced prey rather than crushed sturdy shells.

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Research paper thumbnail of Middle Jurassic (Upper Bathonian and Lower Callovian) jaws of Kosmoceratid ammonites of Central Russia

PalZ, 2020

Elements of the jaw apparatuses of the ammonite genus Kepplerites (Ammonoidea: Stephanoceratoidea... more Elements of the jaw apparatuses of the ammonite genus Kepplerites (Ammonoidea: Stephanoceratoidea, Kosmoceratidae, Keppleritinae) are described from two Upper Bathonian and one Lower Callovian localities of the Russian Platform. The lower jaws (aptychi), based on their size and shape can be assigned to two groups and certainly belonged to the co-occurring macroconchs K. ( Kepplerites ) and theirs microconchs K. ( Toricellites ). It is established that the presence or absence of tuberculate ornamentation on the surface calcite layer in the studied kosmoceratid aptychi (and accordingly the assignment of the aptychi of kosmoceratids to Granulaptychus-type or Praestriaptychus-type) is a result of burial and fossilization in different settings. Most likely all Kosmoceratidae had lower jaws of the Granulaptychus-type, apparently like the related subfamily Garantianinae (family Stephanoceratidae). For the first time, upper jaws of cephalopods supposedly also belonging to the ammonites of genus Kepplerites are described from the Bathonian Stage.

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Research paper thumbnail of Jaw apparatuses of Jurassic Aspidoceratidae (Ammonoidea) and Cope's rule in evolution of aspidoceratid ammonites (comment on “Giant Middle Jurassic (Bathonian) cf. Laevaptychus sp. of the Aztlán section, Hidalgo State, central Mexico” by C.Esquivel-Macías, P.Zell, J.A.Moreno-Bedmar and K.Flores-C...

Journal of South American Earth Sciences, 2021

Comments are provided on a published paper on Middle Jurassic Laevaptychus from central Mexico [C... more Comments are provided on a published paper on Middle Jurassic Laevaptychus from central Mexico [C.EsquivelMacías, P.Zell, J.A.Moreno-Bedmar and K.Flores-Castro, Giant Middle Jurassic (Bathonian) cf. Laevaptychus sp. of the Aztl´ an section, Hidalgo State, central Mexico, Journal of South American Earth Sciences, 110, 103302]. This article describes an interesting finding of large-sized ammonite lower jaws (aptychi referred to Laevaptychus paragenus), claimed as the largest Jurassic aptychi ever known. However, the age of these specimens was erroneously defined due to misidentification of an associated ammonite specimen as Bathonian Procerites. Although poorly preserved, this ammonite shows typical features of the Kimmeridgian genus Idoceras. The Kimmeridgian age of these occurrences is in agreement with findings of Laevaptychus, as this is one of few aptychi formal genera, which belongs to a single ammonite family (Aspidoceratidae). Aspidoceratids appeared in the late Callovian and during the evolution of this lineage maximum sizes of adult specimens and the relative whorl height gradually increased up to Kimmeridgian - Tithonian; only prior to their extinction in early Berriasian, aspidoceratids became uncommon and smaller in size. Laevaptychi are thick-valved aptychi, which have high preservation potential while compared with other aptychi of Jurassic ammonites and their host shells. Giant laevaptychi reported in previous publications (the largest of which reaches 35 cm in length) are briefly reviewed. In adult aspidoceratids the maximum length of aptychi is slightly less than the maximum whorl height. Thus, taking into account the size of the largest aspidoceratid ammonites (up to 85 cm in diameter), the estimated length of the largest laevaptychi can be expected to be ~35–40 cm, which is close to their known record.

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Research paper thumbnail of Early Palaeozoic Discinocarina: a key to the appearance of cephalopod jaws

Lethaia, 2020

Cephalopoda is the only class of molluscs in which virtually all its modern representatives have ... more Cephalopoda is the only class of molluscs in which virtually all its modern representatives have a pair of powerful jaws. There is little doubt that jaws have contributed to the evolutionary success of cephalopods, but their origin still remains a mystery. Though cephalopods appeared at the end of the Cambrian, the oldest unequivocal jaws have been reported to date from the Late Devonian, though they were initially interpreted as phyllopod crustaceans of the suborder Discinocarina. After their relation with ammonoids was proven, they were considered as opercula, and only later their mandibular nature was recognized and widely accepted. Finds of discinocarins from Silurian deposits are still considered as opercula of ammonoid ancestors ‐ nautiloids of the order Orthocerida. However, according to modern ideas, there is no place within their soft body for the location of such large opercula. Moreover, the repeated appearance of very similar structures in the same evolutionary line at least twice, but in different places of the body and for different purposes seems highly improbable. A new hypothesis is proposed herein, in which the Silurian fossils, earlier assigned to Discinocarina, are not specialized opercula, but protective shields, to defend orthocerids not from the predators, but from their own prey. The chitinous plates around the mouth likely appeared in the Silurian orthocerids for protection from such damage and later, during Silurian and Devonian, most likely gradually evolved into the jaws.

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Research paper thumbnail of Recent advances in heteromorph ammonoid palaeobiology

Biological Reviews

Heteromorphs are ammonoids forming a conch with detached whorls (open coiling) or non‐planispiral... more Heteromorphs are ammonoids forming a conch with detached whorls (open coiling) or non‐planispiral coiling. Such aberrant forms appeared convergently four times within this extinct group of cephalopods. Since Wiedmann's seminal paper in this journal, the palaeobiology of heteromorphs has advanced substantially. Combining direct evidence from their fossil record, indirect insights from phylogenetic bracketing, and physical as well as virtual models, we reach an improved understanding of heteromorph ammonoid palaeobiology. Their anatomy, buoyancy, locomotion, predators, diet, palaeoecology, and extinction are discussed. Based on phylogenetic bracketing with nautiloids and coleoids, heteromorphs like other ammonoids had 10 arms, a well‐developed brain, lens eyes, a buccal mass with a radula and a smaller upper as well as a larger lower jaw, and ammonia in their soft tissue. Heteromorphs likely lacked arm suckers, hooks, tentacles, a hood, and an ink sac. All Cretaceous heteromorphs share an aptychus‐type lower jaw with a lamellar calcitic covering. Differences in radular tooth morphology and size in heteromorphs suggest a microphagous diet. Stomach contents of heteromorphs comprise planktic crustaceans, gastropods, and crinoids, suggesting a zooplanktic diet. Forms with a U‐shaped body chamber (ancylocone) are regarded as suspension feeders, whereas orthoconic forms additionally might have consumed benthic prey. Heteromorphs could achieve near‐neutral buoyancy regardless of conch shape or ontogeny. Orthoconic heteromorphs likely had a vertical orientation, whereas ancylocone heteromorphs had a near‐horizontal aperture pointing upwards. Heteromorphs with a U‐shaped body chamber are more stable hydrodynamically than modern Nautilus and were unable substantially to modify their orientation by active locomotion, i.e. they had no or limited access to benthic prey at adulthood. Pathologies reported for heteromorphs were likely inflicted by crustaceans, fish, marine reptiles, and other cephalopods. Pathologies on Ptychoceras corroborates an external shell and rejects the endocochleate hypothesis. Devonian, Triassic, and Jurassic heteromorphs had a preference for deep‐subtidal to offshore facies but are rare in shallow‐subtidal, slope, and bathyal facies. Early Cretaceous heteromorphs preferred deep‐subtidal to bathyal facies. Late Cretaceous heteromorphs are common in shallow‐subtidal to offshore facies. Oxygen isotope data suggest rapid growth and a demersal habitat for adult Discoscaphites and Baculites. A benthic embryonic stage, planktic hatchlings, and a habitat change after one whorl is proposed for Hoploscaphites. Carbon isotope data indicate that some Baculites lived throughout their lives at cold seeps. Adaptation to a planktic life habit potentially drove selection towards smaller hatchlings, implying high fecundity and an ecological role of the hatchlings as micro‐ and mesoplankton. The Chicxulub impact at the Cretaceous/Paleogene (K/Pg) boundary 66 million years ago is the likely trigger for the extinction of ammonoids. Ammonoids likely persisted after this event for 40–500 thousand years and are exclusively represented by heteromorphs. The ammonoid extinction is linked to their small hatchling sizes, planktotrophic diets, and higher metabolic rates than in nautilids, which survived the K/Pg mass extinction event.

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Research paper thumbnail of Discovery of plywood structure in Sphenothallus from Gurovo Formation (Mississippian), Central Russia

Sphenothallus specimens are reported for the first time from the Mississippian of Central Russia.... more Sphenothallus specimens are reported for the first time from the Mississippian of Central Russia. All Sphenothallus specimens have a phosphatic composition and a characteristic laminar structure, which is best observable in the thickened lateral parts of a tube. Most of the lamellae in the tube wall are straight, but some have a wavy morphology and a few are so wrinkled that they form hollow “ribs”. The wrinkled lamellae presumably had an originally higher organic content than the straight lamellae. There are borings on the surfaces of some lamellae that are similar in morphology to the bioerosional traces in various hard, biomineral substrates. Lamellae in the inner parts of the tube wall are composed of fibres. The fibres are parallel to the surface of the tube wall and in successive laminae they differ in orientation by irregularly varying angles. It is possible that the plywood microstructure in Sphenothallus was originally organic and was later phosphatized during fossilization. An alternative, but less likely explanation is that the plywood structure was originally mineralized and therefore is comparable to the phosphatic lamello-fibrillar structures of vertebrates.

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Research paper thumbnail of First teleost otoliths from the Late Jurassic of Russia

Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen, 2020

This is the first record of Late Jurassic (Volgian) otoliths from the boreal province of Russia. ... more This is the first record of Late Jurassic (Volgian) otoliths from the boreal province of Russia. Two new species– Palealbula korchinskyi n. sp. and Palealbula moscoviensis n. sp. – are described representing otoliths of putative stem- albuliform affinities of unresolved familiar position. The phylogeny of Palealbula and the supposedly related genus Protalbula is discussed in the light of the new findings presented in this article. Both genera are shown to represent a common faunal element in the early evolutionary phase of teleosts during the Late Jurassic and Early Cretaceous.

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Research paper thumbnail of Endocerids: suspension feeding nautiloids?

Historical Biology, 2020

For a long time all extinct cephalopods of the subclass Nautiloidea were considered as ecological... more For a long time all extinct cephalopods of the subclass Nautiloidea were considered as ecological analogues of the Recent Nautilus. Recently this view has been rejected: it is now known that among the nautiloids there were not only demersal predators but also epipelagic animals whose life-style and reproduction differed from those of the Nautilus. However, the habits of some nautiloid orders is still poorly understood. One of the most enigmatic cephalopods is the Early Paleozoic nautiloid order Endocerida. Endocerids differ from other nautiloids: they reached gigantic sizes (up to 9 meters), had a wide siphuncle tube and were widespread and numerous during the Ordovician. Since they were an important component of many Ordovician ecosystems, without the understanding of their habits and feeding strategies a correct reconstruction of these ecosystems is impossible. Until now, endocerids have been considered as dominant apex predators, however, this assumption is based on an analogy with the Nautilus mode of life, while the features of the structure of endocerid shells do not confirm this idea and furthermore contradict it. In this article, a new hypothesis is proposed and debated: according to it, the endocerids were planktotrophic cephalopods and the largest of them were giant suspension feeders.

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Research paper thumbnail of A hermit crab preserved inside an ammonite shell from the Upper Jurassic of Central Russia: implications to ammonoid palaeoecology

Palaeogeography, Palaeoclimatology, Palaeoecology, 2020

Abstract The discovery of a hermit crab (superfamily Paguroidea) preserved in the likely immature... more Abstract The discovery of a hermit crab (superfamily Paguroidea) preserved in the likely immature shell of an ammonite, Craspedites nekrassovi is reported from the Upper Jurassic of Moscow, Russia. This is the oldest undoubtable symmetrical hermit crab to date which is known from non-reefal environments. This new occurrence combined with the documentation of numerous sublethal and lethal injuries on ammonite shells in the same beds (probably produced by such paguroids), all suggest that the hermit crabs not only lived in ammonite shells but also hunted these animals. The proportion of damaged shells (including healed ones) varies in different Upper Jurassic ammonite genera from 1.2% in Kachpurites up to 9.3% in Craspedites. Among damaged Kachpurites only 6.25% survived attacks whereas among Craspedites the percentage of survivors was 87.5%. These data imply that Craspedites likely lived near the sea bottom and often encountered hermit crabs, whereas Kachpurites likely lived in the water column.

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Research paper thumbnail of Ammonoid lower jaws of rhynchaptychus type from the Cretaceous of Crimea

Cretaceous Research, 2018

The jaws of Cretaceous nautiloids and some ammonoids (Phylloceratina and Lytoceratina) contain ca... more The jaws of Cretaceous nautiloids and some ammonoids (Phylloceratina and Lytoceratina) contain calcareous elements, namely rhyncholites in the upper jaws and conchorhynchs in the lower. Until now, only several types of numerous rhyncholites have been described from the Cretaceous deposits of Crimea, but lower jaw elements have never been reported from this region. Here we present the first finds of ammonoid lower jaws from the Cenomanian of Crimea that comprise well-preserved calcitic elements. The shape of these conchorhynchs and the ratio of their size to dimensions of the jaw vary in the different specimens. This difference may indicate a variety of food strategies amongst ammonoids with rhynchaptychus-type jaws, whereas all of them were likely durophagous. This assumption is confirmed by longitudinal scratches on the dorsal surface of these conchorhynchs. Amongst dozens of Crimean rhyncholites, specimens which belong to the form genus Tillicheilus have been described previously. Tillicheilus differs fundamentally from other rhyncholites by its shape. A comparison of the conchorhynchs from jaws with Tillicheilus rhyncholites has shown that these calcitic elements are identical, i.e., Tillicheilus constitutes lower jaw elements (conchorhynchs), rather than rhyncholites as interpreted earlier. Finds from Crimea significantly expand the stratigraphical and geographical distribution of Cretaceous rhynchaptychus-type ammonoid jaws.

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Research paper thumbnail of First possible evidence of parasite infestation in Upper Devonian Discosorida (Nautiloidea)

Swiss Journal of Palaeontology

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Research paper thumbnail of Response of cephalopod communities on abrupt environmental changes during the early Aptian OAE1a in the Middle Russian Sea

Cretaceous Research, 2019

Abstract Significant influence of anoxic events on faunal turnovers in marine communities is well... more Abstract Significant influence of anoxic events on faunal turnovers in marine communities is well-established. However, many studies are focused on the impact of anoxic conditions on benthic organisms, while coeval changes in pelagic cephalopod assemblages remain relatively poorly understood. In the present paper we discuss the trends in cephalopod assemblages coinciding with the onset of the early Aptian black shale deposition in European Russia coeval with OAE1a. In few sections of the Saratov Volga area (central part of the Russian Platform), representing both offshore and more proximal nearshore lithofacies of the epicontinental Middle Russian Sea, we have recognized simultaneous changes in ammonite and belemnite successions. Belemnites, represented by the late members of the family Oxyteuthididae, are common in the interval directly preceding anoxic event, but totally disappear with the onset of the black shale deposition. For the Deshayesites ammonites shell size reduction across the mudstone – black shale boundary (maximum shell diameter of adults reduces from ∼20 cm to 7–8 cm) can be observed. Some other ammonites become numerous (Sinzovia) within the black shale interval or show a first occurrence in it (Koeneniceras and Volgoceratoides). In our opinion diminishing of Deshayesites shell size during the early Aptian OAE could be caused by coupling of palaeoenvironmental factors such as progressive warming and regional input of brackish water. Preliminary results of carbon isotope studies of aragonite deriving from the ammonite nacreous layer are also provided.

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Research paper thumbnail of Middle Jurassic ammonoid jaws (anaptychi and rhynchaptychi) from Dagestan, North Caucasus, Russia

Palaeogeography, Palaeoclimatology, Palaeoecology

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Research paper thumbnail of Conellae, enigmatic structures on cephalopod shells – shapes, distribution and formation

Acta Palaeontologica Polonica, 2019

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Research paper thumbnail of Sublethal injuries and abnormalities on embryonic shells of Recent and Jurassic Nautilida

Sublethal injuries and abnormalities of shell walls are common in modern and fossil cephalopods. ... more Sublethal injuries and abnormalities of shell walls are common in modern and fossil cephalopods. In ancient and Recent Nautilida, different abnormalities can even be found on embryonic shells, formed before the hatching. The previously described findings of abnormalities in nautiluses are grooves on the surface of the embryonic shell, as well as curvatures of the shell wall and growth lines. All these abnormalities are related, apparently, with the pressure of the egg membranes or rough surface, on which the egg was attached, and probably with the movement of the embryo within the egg. Sublethal injuries described herein were found on the embryonic shells of Recent and Middle Jurassic (Upper Callovian) nautilids. They include not only the curvature of the shell wall, similar to those described previously, but also rough damage of the apertural edge which arose a long time before hatching. In one case the damaged area is the starting point of a long "scar", indicating the damage of the mantle edge of the embryo. The "scar" is clearly visible on the postembryonic part of the shell and continues almost to the terminal aperture. Such damage of the nautilid embryonic shells has never been described previously. Therefore, it is clear that the range of possible injuries at the embryonic stage of Nautilida is wider than was previously believed and some of these lesions may be associated with an external mechanical action on the egg.

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Research paper thumbnail of First direct evidence of ammonoid ovoviviparity

Ammonoids had high evolutionary rates and diversity throughout their entire history and played an... more Ammonoids had high evolutionary rates and diversity throughout their entire history and played an important role in the high-resolution subdivision 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.

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Research paper thumbnail of Soft-tissue preservation in the Middle Jurassic ammonite Cadoceras from Central Russia

The findings of fossilized ammonite soft tissues are extremely rare, so each specimen may be impo... more The findings of fossilized ammonite soft tissues are extremely rare, so each specimen may be important for understanding the anatomy of these cephalopods. This paper deals with soft tissue fragments and imprints preserved in the rear part of the body chamber of the Middle Jurassic ammonite Cadoceras stupachenkoi (Mitta) from Central Russia. At the base of the body chamber of this ammonite in front of the last septum, a mantle fragment with clearly visible longitudinal fibers and imprints of the palliovisceral ligament are preserved. In front and slightly to the side of the mantle fragment, a small area with branched structures is located; probably, these structures are fragments of gills.
In general, the structure of the soft tissues in the rear part of the ammonite body looks very similar to that of modern nautilids, with one exception: mantle fibers are not directed forward as observed in Nautilus, but to the mid-ventral line, probably to the ventral muscle.

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Research paper thumbnail of The first record of jaws of Boreal Valanginian ammonites (Cephalopoda, Polyptychitidae

Jaws of ammonites which inhabited the Panboreal Superrealm during the Jurassic and Cretaceous are... more Jaws of ammonites which inhabited the Panboreal Superrealm during the Jurassic and Cretaceous are poorly known in comparison to those of Tethyan ammonoid faunas. This paucity may be explained by limited thickness, or even absence of an outer calcitic layer, in lower jaw elements (aptychi) of Boreal ammonites. Here we describe, for the first time, the jaws (both lower and upper) of ammonites of the Boreal family Polyptychitidae, of Early Cretaceous (Valanginian) age. Polyptychitid lower jaws are of the aptychus type, but have an unusual pointed and convex shape. However, lower jaws of Late Jurassic ancestors of polyptychitids (Craspeditidae) as well as Middle Jurassic cardioceratids (Pseudocadoceras) have a near-identical shape, as do previously described aptychi of the Late Cretaceous genera Neogastroplites and Placenticeras (Hoplitoidea). The close resemblance of lower jaws of evolutionarily distant ammonites may be linked to a similar lifestyle, but more data are needed to substantiate this. Upper jaws of polyptychitid are closely similar to previously described upper jaws of Jurassic ammonites, which indicates the conservatism of this part of the jaw apparatus. Together with shells and jaws of the Valanginian ammonites described herein, jaws of coleoids (likely belemnites) as well as arm hooks (onychites) have been found.

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Research paper thumbnail of First discovery of the soft‐body imprint of an Oligocene fossil squid indicates its piscivorous diet

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Research paper thumbnail of Structure of the calcitic layer of the aptychus of the ammonite genus Peltoceras

Annales Societatis Geologorum Poloniae, 2021

The aptychi of ammonites combined the functions of lower jaws and protective opercula. They consi... more The aptychi of ammonites combined the functions of lower jaws and protective opercula. They consist of two parts: an inner organic layer and an outer calcitic lamella. In different evolutionary lineages of ammonites, the shape of aptychi, the sculpture of their surface and the microstructure of the calcitic layer vary greatly. However, the structure of the aptychi is not known for all evolutionary lineages of ammonites. Although numerous aptychi have been described for the Jurassic family Aspidoceratidae, almost all of them belong to only one evolutionary branch of this family – the Aspidoceratinae (sensu lato). For the second branch – the Peltoceratinae, only one aptychus had been described to date and the structure of its calcitic layer remained unknown. In this article, for the first time, the structure of the aptychus of the Peltoceratinae (upper Callovian Peltoceras) is described. The surface of this aptychus is covered with rough ribs and the calcitic part consists of only one layer of dense calcite. The thickness of the aptychus is much greater than that of the aptychi of supposed ancestors of the Peltoceratinae. The increase in the thickness of the aptychi in both the Aspidoceratinae and the Peltoceratinae, contemporaneously with the appearance of spines on their shells, is most likely related to increasing the protective function of the aptychi of these ammonites in the late Callovian.

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Research paper thumbnail of An unusual conchorhynch from the upper Maastrichtian of the southeast Netherlands and the distinction between nautiloid and ammonoid conchorhynchs (Mollusca, Cephalopoda)⁋

Cretaceous Research, 2022

A single, atypical conchorhynch (calcitic tip of a cephalopod lower jaw), recovered from the uppe... more A single, atypical conchorhynch (calcitic tip of a cephalopod lower jaw), recovered from the uppermost Meerssen Member (Maastricht Formation, upper Maastrichtian) at the former ENCI-HeidelbergCement Group quarry, south of Maastricht, is described as a new parataxon, Conchorhynchus illustris sp. nov. The specimen can be differentiated from all previous conchorhynch records on account of its large size, elongated shape and, in particular, of the structure of its apical part which is smooth and forwardly elongated. During the Late Cretaceous, conchorhynchs formed part of the jaw apparatus of nautilids and of two ammonoid suborders, Phylloceratina and Lytoceratina. Since conchorhynchs are most often found separated from jaws, establishing to which group of cephalopods their bearer belonged can be complicated. Here, for the first time, we propose a set of morphological criteria to differentiate clearly between nautiloid and ammonoid conchorhynchs. Although Conchorhynchus illustris sp. nov. is distinct from all currently known nautilid conchorhynchs, the sum of its morphological features is indicative of assignment to that cephalopod group. The upper portion of the Maastricht Formation in the Maastricht area (Nekum and Meerssen members) has yielded internal and external moulds of shells of the nautilid Eutrephoceras and the hercoglossid Cimomia. The new conchorhynch type described herein most likely belonged to one of these shell-based taxa. Judging from its unusual shape, the feeding strategy of its bearer must have differed from that of modern nautilids, in that it held and pierced prey rather than crushed sturdy shells.

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Research paper thumbnail of Middle Jurassic (Upper Bathonian and Lower Callovian) jaws of Kosmoceratid ammonites of Central Russia

PalZ, 2020

Elements of the jaw apparatuses of the ammonite genus Kepplerites (Ammonoidea: Stephanoceratoidea... more Elements of the jaw apparatuses of the ammonite genus Kepplerites (Ammonoidea: Stephanoceratoidea, Kosmoceratidae, Keppleritinae) are described from two Upper Bathonian and one Lower Callovian localities of the Russian Platform. The lower jaws (aptychi), based on their size and shape can be assigned to two groups and certainly belonged to the co-occurring macroconchs K. ( Kepplerites ) and theirs microconchs K. ( Toricellites ). It is established that the presence or absence of tuberculate ornamentation on the surface calcite layer in the studied kosmoceratid aptychi (and accordingly the assignment of the aptychi of kosmoceratids to Granulaptychus-type or Praestriaptychus-type) is a result of burial and fossilization in different settings. Most likely all Kosmoceratidae had lower jaws of the Granulaptychus-type, apparently like the related subfamily Garantianinae (family Stephanoceratidae). For the first time, upper jaws of cephalopods supposedly also belonging to the ammonites of genus Kepplerites are described from the Bathonian Stage.

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Research paper thumbnail of Jaw apparatuses of Jurassic Aspidoceratidae (Ammonoidea) and Cope's rule in evolution of aspidoceratid ammonites (comment on “Giant Middle Jurassic (Bathonian) cf. Laevaptychus sp. of the Aztlán section, Hidalgo State, central Mexico” by C.Esquivel-Macías, P.Zell, J.A.Moreno-Bedmar and K.Flores-C...

Journal of South American Earth Sciences, 2021

Comments are provided on a published paper on Middle Jurassic Laevaptychus from central Mexico [C... more Comments are provided on a published paper on Middle Jurassic Laevaptychus from central Mexico [C.EsquivelMacías, P.Zell, J.A.Moreno-Bedmar and K.Flores-Castro, Giant Middle Jurassic (Bathonian) cf. Laevaptychus sp. of the Aztl´ an section, Hidalgo State, central Mexico, Journal of South American Earth Sciences, 110, 103302]. This article describes an interesting finding of large-sized ammonite lower jaws (aptychi referred to Laevaptychus paragenus), claimed as the largest Jurassic aptychi ever known. However, the age of these specimens was erroneously defined due to misidentification of an associated ammonite specimen as Bathonian Procerites. Although poorly preserved, this ammonite shows typical features of the Kimmeridgian genus Idoceras. The Kimmeridgian age of these occurrences is in agreement with findings of Laevaptychus, as this is one of few aptychi formal genera, which belongs to a single ammonite family (Aspidoceratidae). Aspidoceratids appeared in the late Callovian and during the evolution of this lineage maximum sizes of adult specimens and the relative whorl height gradually increased up to Kimmeridgian - Tithonian; only prior to their extinction in early Berriasian, aspidoceratids became uncommon and smaller in size. Laevaptychi are thick-valved aptychi, which have high preservation potential while compared with other aptychi of Jurassic ammonites and their host shells. Giant laevaptychi reported in previous publications (the largest of which reaches 35 cm in length) are briefly reviewed. In adult aspidoceratids the maximum length of aptychi is slightly less than the maximum whorl height. Thus, taking into account the size of the largest aspidoceratid ammonites (up to 85 cm in diameter), the estimated length of the largest laevaptychi can be expected to be ~35–40 cm, which is close to their known record.

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Research paper thumbnail of Early Palaeozoic Discinocarina: a key to the appearance of cephalopod jaws

Lethaia, 2020

Cephalopoda is the only class of molluscs in which virtually all its modern representatives have ... more Cephalopoda is the only class of molluscs in which virtually all its modern representatives have a pair of powerful jaws. There is little doubt that jaws have contributed to the evolutionary success of cephalopods, but their origin still remains a mystery. Though cephalopods appeared at the end of the Cambrian, the oldest unequivocal jaws have been reported to date from the Late Devonian, though they were initially interpreted as phyllopod crustaceans of the suborder Discinocarina. After their relation with ammonoids was proven, they were considered as opercula, and only later their mandibular nature was recognized and widely accepted. Finds of discinocarins from Silurian deposits are still considered as opercula of ammonoid ancestors ‐ nautiloids of the order Orthocerida. However, according to modern ideas, there is no place within their soft body for the location of such large opercula. Moreover, the repeated appearance of very similar structures in the same evolutionary line at least twice, but in different places of the body and for different purposes seems highly improbable. A new hypothesis is proposed herein, in which the Silurian fossils, earlier assigned to Discinocarina, are not specialized opercula, but protective shields, to defend orthocerids not from the predators, but from their own prey. The chitinous plates around the mouth likely appeared in the Silurian orthocerids for protection from such damage and later, during Silurian and Devonian, most likely gradually evolved into the jaws.

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Research paper thumbnail of Recent advances in heteromorph ammonoid palaeobiology

Biological Reviews

Heteromorphs are ammonoids forming a conch with detached whorls (open coiling) or non‐planispiral... more Heteromorphs are ammonoids forming a conch with detached whorls (open coiling) or non‐planispiral coiling. Such aberrant forms appeared convergently four times within this extinct group of cephalopods. Since Wiedmann's seminal paper in this journal, the palaeobiology of heteromorphs has advanced substantially. Combining direct evidence from their fossil record, indirect insights from phylogenetic bracketing, and physical as well as virtual models, we reach an improved understanding of heteromorph ammonoid palaeobiology. Their anatomy, buoyancy, locomotion, predators, diet, palaeoecology, and extinction are discussed. Based on phylogenetic bracketing with nautiloids and coleoids, heteromorphs like other ammonoids had 10 arms, a well‐developed brain, lens eyes, a buccal mass with a radula and a smaller upper as well as a larger lower jaw, and ammonia in their soft tissue. Heteromorphs likely lacked arm suckers, hooks, tentacles, a hood, and an ink sac. All Cretaceous heteromorphs share an aptychus‐type lower jaw with a lamellar calcitic covering. Differences in radular tooth morphology and size in heteromorphs suggest a microphagous diet. Stomach contents of heteromorphs comprise planktic crustaceans, gastropods, and crinoids, suggesting a zooplanktic diet. Forms with a U‐shaped body chamber (ancylocone) are regarded as suspension feeders, whereas orthoconic forms additionally might have consumed benthic prey. Heteromorphs could achieve near‐neutral buoyancy regardless of conch shape or ontogeny. Orthoconic heteromorphs likely had a vertical orientation, whereas ancylocone heteromorphs had a near‐horizontal aperture pointing upwards. Heteromorphs with a U‐shaped body chamber are more stable hydrodynamically than modern Nautilus and were unable substantially to modify their orientation by active locomotion, i.e. they had no or limited access to benthic prey at adulthood. Pathologies reported for heteromorphs were likely inflicted by crustaceans, fish, marine reptiles, and other cephalopods. Pathologies on Ptychoceras corroborates an external shell and rejects the endocochleate hypothesis. Devonian, Triassic, and Jurassic heteromorphs had a preference for deep‐subtidal to offshore facies but are rare in shallow‐subtidal, slope, and bathyal facies. Early Cretaceous heteromorphs preferred deep‐subtidal to bathyal facies. Late Cretaceous heteromorphs are common in shallow‐subtidal to offshore facies. Oxygen isotope data suggest rapid growth and a demersal habitat for adult Discoscaphites and Baculites. A benthic embryonic stage, planktic hatchlings, and a habitat change after one whorl is proposed for Hoploscaphites. Carbon isotope data indicate that some Baculites lived throughout their lives at cold seeps. Adaptation to a planktic life habit potentially drove selection towards smaller hatchlings, implying high fecundity and an ecological role of the hatchlings as micro‐ and mesoplankton. The Chicxulub impact at the Cretaceous/Paleogene (K/Pg) boundary 66 million years ago is the likely trigger for the extinction of ammonoids. Ammonoids likely persisted after this event for 40–500 thousand years and are exclusively represented by heteromorphs. The ammonoid extinction is linked to their small hatchling sizes, planktotrophic diets, and higher metabolic rates than in nautilids, which survived the K/Pg mass extinction event.

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Research paper thumbnail of Discovery of plywood structure in Sphenothallus from Gurovo Formation (Mississippian), Central Russia

Sphenothallus specimens are reported for the first time from the Mississippian of Central Russia.... more Sphenothallus specimens are reported for the first time from the Mississippian of Central Russia. All Sphenothallus specimens have a phosphatic composition and a characteristic laminar structure, which is best observable in the thickened lateral parts of a tube. Most of the lamellae in the tube wall are straight, but some have a wavy morphology and a few are so wrinkled that they form hollow “ribs”. The wrinkled lamellae presumably had an originally higher organic content than the straight lamellae. There are borings on the surfaces of some lamellae that are similar in morphology to the bioerosional traces in various hard, biomineral substrates. Lamellae in the inner parts of the tube wall are composed of fibres. The fibres are parallel to the surface of the tube wall and in successive laminae they differ in orientation by irregularly varying angles. It is possible that the plywood microstructure in Sphenothallus was originally organic and was later phosphatized during fossilization. An alternative, but less likely explanation is that the plywood structure was originally mineralized and therefore is comparable to the phosphatic lamello-fibrillar structures of vertebrates.

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Research paper thumbnail of First teleost otoliths from the Late Jurassic of Russia

Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen, 2020

This is the first record of Late Jurassic (Volgian) otoliths from the boreal province of Russia. ... more This is the first record of Late Jurassic (Volgian) otoliths from the boreal province of Russia. Two new species– Palealbula korchinskyi n. sp. and Palealbula moscoviensis n. sp. – are described representing otoliths of putative stem- albuliform affinities of unresolved familiar position. The phylogeny of Palealbula and the supposedly related genus Protalbula is discussed in the light of the new findings presented in this article. Both genera are shown to represent a common faunal element in the early evolutionary phase of teleosts during the Late Jurassic and Early Cretaceous.

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Research paper thumbnail of Endocerids: suspension feeding nautiloids?

Historical Biology, 2020

For a long time all extinct cephalopods of the subclass Nautiloidea were considered as ecological... more For a long time all extinct cephalopods of the subclass Nautiloidea were considered as ecological analogues of the Recent Nautilus. Recently this view has been rejected: it is now known that among the nautiloids there were not only demersal predators but also epipelagic animals whose life-style and reproduction differed from those of the Nautilus. However, the habits of some nautiloid orders is still poorly understood. One of the most enigmatic cephalopods is the Early Paleozoic nautiloid order Endocerida. Endocerids differ from other nautiloids: they reached gigantic sizes (up to 9 meters), had a wide siphuncle tube and were widespread and numerous during the Ordovician. Since they were an important component of many Ordovician ecosystems, without the understanding of their habits and feeding strategies a correct reconstruction of these ecosystems is impossible. Until now, endocerids have been considered as dominant apex predators, however, this assumption is based on an analogy with the Nautilus mode of life, while the features of the structure of endocerid shells do not confirm this idea and furthermore contradict it. In this article, a new hypothesis is proposed and debated: according to it, the endocerids were planktotrophic cephalopods and the largest of them were giant suspension feeders.

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Research paper thumbnail of A hermit crab preserved inside an ammonite shell from the Upper Jurassic of Central Russia: implications to ammonoid palaeoecology

Palaeogeography, Palaeoclimatology, Palaeoecology, 2020

Abstract The discovery of a hermit crab (superfamily Paguroidea) preserved in the likely immature... more Abstract The discovery of a hermit crab (superfamily Paguroidea) preserved in the likely immature shell of an ammonite, Craspedites nekrassovi is reported from the Upper Jurassic of Moscow, Russia. This is the oldest undoubtable symmetrical hermit crab to date which is known from non-reefal environments. This new occurrence combined with the documentation of numerous sublethal and lethal injuries on ammonite shells in the same beds (probably produced by such paguroids), all suggest that the hermit crabs not only lived in ammonite shells but also hunted these animals. The proportion of damaged shells (including healed ones) varies in different Upper Jurassic ammonite genera from 1.2% in Kachpurites up to 9.3% in Craspedites. Among damaged Kachpurites only 6.25% survived attacks whereas among Craspedites the percentage of survivors was 87.5%. These data imply that Craspedites likely lived near the sea bottom and often encountered hermit crabs, whereas Kachpurites likely lived in the water column.

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Research paper thumbnail of Ammonoid lower jaws of rhynchaptychus type from the Cretaceous of Crimea

Cretaceous Research, 2018

The jaws of Cretaceous nautiloids and some ammonoids (Phylloceratina and Lytoceratina) contain ca... more The jaws of Cretaceous nautiloids and some ammonoids (Phylloceratina and Lytoceratina) contain calcareous elements, namely rhyncholites in the upper jaws and conchorhynchs in the lower. Until now, only several types of numerous rhyncholites have been described from the Cretaceous deposits of Crimea, but lower jaw elements have never been reported from this region. Here we present the first finds of ammonoid lower jaws from the Cenomanian of Crimea that comprise well-preserved calcitic elements. The shape of these conchorhynchs and the ratio of their size to dimensions of the jaw vary in the different specimens. This difference may indicate a variety of food strategies amongst ammonoids with rhynchaptychus-type jaws, whereas all of them were likely durophagous. This assumption is confirmed by longitudinal scratches on the dorsal surface of these conchorhynchs. Amongst dozens of Crimean rhyncholites, specimens which belong to the form genus Tillicheilus have been described previously. Tillicheilus differs fundamentally from other rhyncholites by its shape. A comparison of the conchorhynchs from jaws with Tillicheilus rhyncholites has shown that these calcitic elements are identical, i.e., Tillicheilus constitutes lower jaw elements (conchorhynchs), rather than rhyncholites as interpreted earlier. Finds from Crimea significantly expand the stratigraphical and geographical distribution of Cretaceous rhynchaptychus-type ammonoid jaws.

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Research paper thumbnail of First possible evidence of parasite infestation in Upper Devonian Discosorida (Nautiloidea)

Swiss Journal of Palaeontology

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Research paper thumbnail of Response of cephalopod communities on abrupt environmental changes during the early Aptian OAE1a in the Middle Russian Sea

Cretaceous Research, 2019

Abstract Significant influence of anoxic events on faunal turnovers in marine communities is well... more Abstract Significant influence of anoxic events on faunal turnovers in marine communities is well-established. However, many studies are focused on the impact of anoxic conditions on benthic organisms, while coeval changes in pelagic cephalopod assemblages remain relatively poorly understood. In the present paper we discuss the trends in cephalopod assemblages coinciding with the onset of the early Aptian black shale deposition in European Russia coeval with OAE1a. In few sections of the Saratov Volga area (central part of the Russian Platform), representing both offshore and more proximal nearshore lithofacies of the epicontinental Middle Russian Sea, we have recognized simultaneous changes in ammonite and belemnite successions. Belemnites, represented by the late members of the family Oxyteuthididae, are common in the interval directly preceding anoxic event, but totally disappear with the onset of the black shale deposition. For the Deshayesites ammonites shell size reduction across the mudstone – black shale boundary (maximum shell diameter of adults reduces from ∼20 cm to 7–8 cm) can be observed. Some other ammonites become numerous (Sinzovia) within the black shale interval or show a first occurrence in it (Koeneniceras and Volgoceratoides). In our opinion diminishing of Deshayesites shell size during the early Aptian OAE could be caused by coupling of palaeoenvironmental factors such as progressive warming and regional input of brackish water. Preliminary results of carbon isotope studies of aragonite deriving from the ammonite nacreous layer are also provided.

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Research paper thumbnail of Middle Jurassic ammonoid jaws (anaptychi and rhynchaptychi) from Dagestan, North Caucasus, Russia

Palaeogeography, Palaeoclimatology, Palaeoecology

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Research paper thumbnail of Conellae, enigmatic structures on cephalopod shells – shapes, distribution and formation

Acta Palaeontologica Polonica, 2019

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Research paper thumbnail of Sublethal injuries and abnormalities on embryonic shells of Recent and Jurassic Nautilida

Sublethal injuries and abnormalities of shell walls are common in modern and fossil cephalopods. ... more Sublethal injuries and abnormalities of shell walls are common in modern and fossil cephalopods. In ancient and Recent Nautilida, different abnormalities can even be found on embryonic shells, formed before the hatching. The previously described findings of abnormalities in nautiluses are grooves on the surface of the embryonic shell, as well as curvatures of the shell wall and growth lines. All these abnormalities are related, apparently, with the pressure of the egg membranes or rough surface, on which the egg was attached, and probably with the movement of the embryo within the egg. Sublethal injuries described herein were found on the embryonic shells of Recent and Middle Jurassic (Upper Callovian) nautilids. They include not only the curvature of the shell wall, similar to those described previously, but also rough damage of the apertural edge which arose a long time before hatching. In one case the damaged area is the starting point of a long "scar", indicating the damage of the mantle edge of the embryo. The "scar" is clearly visible on the postembryonic part of the shell and continues almost to the terminal aperture. Such damage of the nautilid embryonic shells has never been described previously. Therefore, it is clear that the range of possible injuries at the embryonic stage of Nautilida is wider than was previously believed and some of these lesions may be associated with an external mechanical action on the egg.

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Research paper thumbnail of First direct evidence of ammonoid ovoviviparity

Ammonoids had high evolutionary rates and diversity throughout their entire history and played an... more Ammonoids had high evolutionary rates and diversity throughout their entire history and played an important role in the high-resolution subdivision 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.

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Research paper thumbnail of Soft-tissue preservation in the Middle Jurassic ammonite Cadoceras from Central Russia

The findings of fossilized ammonite soft tissues are extremely rare, so each specimen may be impo... more The findings of fossilized ammonite soft tissues are extremely rare, so each specimen may be important for understanding the anatomy of these cephalopods. This paper deals with soft tissue fragments and imprints preserved in the rear part of the body chamber of the Middle Jurassic ammonite Cadoceras stupachenkoi (Mitta) from Central Russia. At the base of the body chamber of this ammonite in front of the last septum, a mantle fragment with clearly visible longitudinal fibers and imprints of the palliovisceral ligament are preserved. In front and slightly to the side of the mantle fragment, a small area with branched structures is located; probably, these structures are fragments of gills.
In general, the structure of the soft tissues in the rear part of the ammonite body looks very similar to that of modern nautilids, with one exception: mantle fibers are not directed forward as observed in Nautilus, but to the mid-ventral line, probably to the ventral muscle.

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Research paper thumbnail of Early Palaeozoic Discinocarina and a new hypothesis of the appearance of cephalopod jaws

The jaws of Cephalopoda are a unique structure among molluscs; however, their origin still remain... more The jaws of Cephalopoda are a unique structure among molluscs; however, their origin still remains a mystery. Whereas cephalopods appeared at the end of the Cambrian, their oldest jaws have been reported to date from the Upper Devonian, and belonged to ammonoids. However, in the Silurian and Devonian strata, initially organic structures related to cephalopods can be found. The most widely known among them is Aptychopsis, several specimens of which were found in the apertures of orthocerid shells. These structures had initially been interpreted as phyllopod crustaceans (suborder Discinocarina), but now their Devonian representatives are thought to be the lower jaws of ammonoids. ilurian discinocarins, including Aptychopsis, are considered as opercula of ammonoid ancestors – orthocerid nautiloids. However, it is unlikely that very similar structures appeared in the same evolutionary lineage twice, but in different body parts and for different purposes. It seems more possible that the Silurian discinocarins are basal proto-jaws. Here I propose a new hypothesis, according to which initially organic plates arose on the front of the nautiloid head around the radula to protect them not from predators, but from resisting prey. They had some mobility to release the radula and gradually evolved into a jaw apparatus.

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