Three previously unknown octopods are described from Upper Cenomanian limestones of the Hâqel and Hâdjoula localities (Lebanon). Keuppia levante gen. nov., sp. nov., Keuppia hyperbolaris gen. nov,. sp. nov. and Styletoctopus annae gen. nov, . sp. nov. are regarded as the earliest representatives of the Octopoda (= Incirrata). This assumption is mainly based on their medially isolated bipartite gladius vestige. As can be inferred from growth increments, Keuppia gen. nov. can be distinguished from the genus Palaeoctopus by blades that grow forwards along their longitudinal axis. The gladius vestige of Keuppia hyperbolaris sp. nov. differs from that of Keuppia levante sp. nov. in having a more heterogeneous course of growth lines. Based on a pair of widely separated stylets, which closely resemble the rods of modern octopods, Styletoctopus annae gen. nov., sp. nov. is assigned to the Recent family Octopodidae. Peculiar encrustations, which are situated in close association with the gladius vestiges of Keuppia levante sp. nov., Keuppia hyperbolaris sp. nov., and Styletoctopus annae sp. nov. are interpreted as basal fin cartilages. The gladius vestige morphology of Keuppia hyperbolaris sp. nov. and Keuppia levante sp. nov. opens the possibility that both the Octopda and the Cirroctopoda originated from loligosepiid vampyropods instead of teudopseid. The surprising existence of a stylet‐like gladius vestige in Styletoctopus annae sp. nov. suggests that the octopod clade branched off much earlier than previously believed. Octopod apomorphies such as the development of stylets, loss of fins and cirri must have been occurred before the Cenomanian.
Body-size reduction is considered an important response to current climate warming and has been observed during past biotic crises, including the Pliensbachian–Toarcian crisis, a second-order mass extinction. However, in fossil cephalopod studies, the mechanisms and their potential link with climate are rarely investigated and palaeobiological scales of organization are not usually differentiated. Here, we hypothesize that belemnites reduce their adult size across the Pliensbachian–Toarcian boundary warming event. Belemnite body-size dynamics across the Pliensbachian–Toarcian boundary in the Peniche section (Lusitanian Basin, Portugal) were analysed based on the newly collected field data. We disentangle the mechanisms and the environmental drivers of the size fluctuations observed from the individual to the assemblage scale. Despite the lack of a major taxonomic turnover, a 40% decrease in rostrum volume is observed across the Pliensbachian–Toarcian boundary, before the Toarcian Oceanic Anoxic Event where belemnites go locally extinct. The pattern is mainly driven by a reduction in adult size of the two dominant species, Pseudohastites longiformis and Passaloteuthis bisulcata. Belemnite-size distribution is best correlated with fluctuations in a palaeotemperature proxy (stable oxygen isotopes); however, potential indirect effects of volcanism and carbon cycle perturbations may also play a role. This highlights the complex interplay between environmental stressors (warming, deoxygenation, nutrient input) and biotic variables (productivity, competition, migration) associated with these hyperthermal events in driving belemnite body-size.
N. 2014: Embryonic shell structure of Early-Middle Jurassic belemnites, and its significance for belemnite expansion and diversification in the Jurassic. Lethaia, Vol. 47, Early Jurassic belemnites are of particular interest to the study of the evolution of skeletal morphology in Lower Carboniferous to the uppermost Cretaceous belemnoids, because they signal the beginning of a global Jurassic-Cretaceous expansion and diversification of belemnitids. We investigated potentially relevant, to this evolutionary pattern, shell features of Sinemurian-Bajocian Nannobelus, Parapassaloteuthis, Holcobelus and Pachybelemnopsis from the Paris Basin. Our analysis of morphological, ultrastructural and chemical traits of the earliest ontogenetic stages of the shell suggests that modified embryonic shell structure of Early-Middle Jurassic belemnites was a factor in their expansion and colonization of the pelagic zone and resulted in remarkable diversification of belemnites. Innovative traits of the embryonic shell of SinemurianBajocian belemnites include: (1) an inorganic-organic primordial rostrum encapsulating the protoconch and the phragmocone, its non-biomineralized component, possibly chitin, is herein detected for the first time; (2) an organic rich closing membrane which was under formation. It was yet perforated and possessed a foramen; and (3) an organic rich pro-ostracum earlier documented in an embryonic shell of Pliensbachian Passaloteuthis. The inorganic-organic primordial rostrum tightly coating the protoconch and phragmocone supposedly enhanced protection, without increase in shell weight, of the Early Jurassic belemnites against explosion in deepwater environment. This may have increased the depth and temperature ranges of hatching eggs, accelerated the adaptation of hatchlings to a nektonic mode of life and promoted increasing diversity of belemnoids. This study supports the hypothesis that belemnite hatchlings were 'a miniature of the adults'. □ Belemnites, embryonic shell, expansion, inorganic-organic shell matter, Jurassic.
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