LATE CAMBRIAN PLECTRONOCERID NAUTILOIDS AND THEIR ROLE IN CEPHALOPOD EVOLUTION

Abstract:  Numerous plectronocerid nautiloids appear in the Upper Cambrian of China. We have restudied their siphuncular structure, first described some 20 years ago. The siphuncle is characterized by: (1) long and holochoanitic septal necks dorsally but short and recurved necks laterally and ventrally; (2) strongly expanded connecting rings laterally; (3) two calcified layers in each connecting ring, outer spherulitic-prismatic and inner compact, the latter perforated by numerous pore canals; and (4) highly oblique siphuncular segments. The strongly expanded lateral sides of the connecting rings, together with the highly oblique course of the siphuncular segments, considerably enlarged the surface area of the connecting rings in each chamber, thereby increasing the transport capacity of cameral liquid. Thus, from their first appearance, plectronocerid nautiloids had developed a siphuncle for the replacement of cameral liquid with gases, and this system had a better and a more sophisticated design than that seen in stratigraphically younger nautiloids. However, their small orthoconic or slightly cyrtoconic shells were not well adapted for jet-powered swimming.     

Siphuncular structure in the orders Tarphycerida and Barrandeocerida (Cephalopoda: Nautiloidea)

Abstract: The siphuncular structure is described in two Silurian taxa, Boionautilus tyrannus and Cumingsoceras complanatus, currently placed in the Tarpycerida. Tarphycerids have the Nautilus type of connecting ring that is composed of an outer, thick, spherulitic‐prismatic layer and an inner glycoprotein layer, the latter was destroyed by diagenesis. However, both Silurian specimens have the connecting ring of the calcified‐perforate type, previously known to occur in orthocerids, actinocerids, plectronocerids and now also in barrandeocerids. In this type, the inner layer of the connecting ring is calcified and perforated by pore canals. Boionautilus and Cumingsoceras are therefore classified with barrandeocerids and not with tarphycerids.     

Endocochliate embryo model in the Mesozoic Ammonitida

An endocochliate embryo model for the Mesozoic Ammonitida is proposed based on scanning electron microscopy of the ammonitella (= embryonic shell) stage of well‐preserved Japanese Cretaceous specimens belonging to nine species of five superfamilies. As in other specimens described previously, the ammonitella wall succeeding from the initial chamber ("protoconch") in the species examined consists of the inner prismatic, middle subprismatic and outer prismatic layers, with minute tubercles resting on the outer. Developmental patterns of these structures and their comparison with primary shell formation in modern Nautilus and Spirula suggest that the outer thin prismatic layer with microtubercles was secreted by the exterior epithelium after the completion of the main ammonitella wall by the interior shell gland. Thus, the early ammonite embryo might have had an endocochliate structural plan as in coleoids, and at the time of hatching the overlying mantle epithelium had shifted anteriorly to become an ectocochliate condition.     

Function of calcium phosphate renal concrements in extant Nautilus: a paradigm for Cambrian-relict short-term mineral reserve equivalent to vertebrate bone

Renal uroliths (concrements) of calcium phosphate have long been known to exist in both growing and mature (non-growing) Nautilus specimens, but to date no evidence-based explanation for their existence has been available. The currently favored speculation is that they function as a calcium reserve for shell and septal calcification. Here we present new observational and experimental data that are consistent with the hypothesis that they serve as a mineral/ion reserve, allowing short-term (<1 day) addition of ionized calcium and phosphorus to blood and other body fluids, in a way analogous to that of vertebrate bone. In both in-ocean experiments and during long-term observation of captive nautiluses, concrements disappear during two different, energy-intensive activities involving removal of anions and cations from newly secreted cameral liquid in the chamber formation cycle, and during dives to depths requiring high osmotic pressures within the canaliculi of the siphuncular epithelium to keep previously emptied chambers from flooding due to suddenly increased ambient hydrostatic pressure. New concrements reappear at other points in the chamber formation cycle and when normal living depth is restored. The use of concrements as an ion reserve and the Cambrian ancestry of nautiloids indicate that Nautilus may exemplify a solution to the problem of energy supply in newly evolved swimmers of the Cambrian radiation independent of that seen in fish.     

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.     

Characterization of actinoceratoid cephalopods by their siphuncular structure

Partially phosphatized connecting rings show well-preserved structural details in the actinoceratoid Adamsoceras holmi (Ormoceratidae) from the Lower Ordovician (Kundan) of Estonia. Each connecting ring is composed of an outer, thin, spherulitic-prismatic layer, and an inner, thick, calcareous, lamellar layer, the latter being traversed by numerous large pores. The calcareous lamellar inner layer occurs in the connecting rings of three other actinoceratoids: Eushan-tungoceras pseudoimbricatum, Rayonnoceras solidiforme , and Huroniella sp. In Huroniella sp. and R. solidiforme , this layer is traversed by similar large pores as in Adamsoceras holmi , but in E. pseudoimbricatum the pores were probably too narrow to be recognized. The described structural type of the connecting rings, together with well-developed annular endosiphuncular deposits, are here considered as characteristic for actinoceratoids. The inflated connecting ring in actinoceratoids had a great surface area, which increased the number of pores across it. The permeability and emptying rates of the cameral liquid through the connecting ring could therefore have been as high or higher than that in Nautilus . In being composed of numerous calcified lamellae, the mechanical strength of the connecting ring against hydrostatic pressure could have also been the same or higher than that in Nautilus.     

New insights into the buccal apparatus of the Goniatitina: palaeobiological and phylogenetic implications

Exceptionally well‐preserved radulae and lower jaws found both inside the body chamber of Cravenoceras fayettevillae (Mississippian) and in isolated nodules (Mississippian and Pennsylvanian) were analysed using CT scan and synchrotron propagation phase‐contrast tomography. This approach reveals new anatomical details allowing us to investigate wear, preservation and muscle insertion in the buccal mass of the Goniatitina. For comparison, Recent cephalopod radulae were also investigated. The radula in the Goniatitina consists of nine elements per row, and its morphology is conservative in the group. The shape of the teeth is similar to that in some Recent coleoids. The lower jaw is morphologically closer to that of Recent Nautilus, consisting of a wide outer lamella and a reduced inner lamella. The morphology of the inner lamella reflects the outer lamella in smaller dimensions and does not protrude posteroventrally. The space between the inner and outer lamellae allows for muscle insertion. Our morphological data indicate that a mosaic of characters is present in the buccal mass of Goniatitina with some parts of the buccal mass being more primitive than others. This implies that different parts of the buccal mass may have followed different evolutionary histories.     

Taxonomy,morphology and phylogeny of Late Cretaceous spirulid coleoids (Cephalopoda) from Greenland and Canada

Abstract: Groenlandibelus rosenkrantzi from the Maastrichtian of Greenland has long been thought to constitute an early representative of spirulid coleoids. This study shows that this view must be reassessed, at least in part. A re‐investigation of the types and of material recorded subsequently has revealed that none of these specimens is conspecific with the holotype of G. rosenkrantzi. Cyrtobelus birkelundae gen. nov, sp. nov. differs from the type of G. rosenkrantzi in having lower chambers and in lacking an apically elongated sheath. The longiconic phragmocone of G. rosenkrantzi has more features in common with the presumed spirulid genus Naefia. A specimen described in detail by J. A. Jeletzky in the mid 1960s as ‘G. rosenkrantzi’ is designated holotype of C. birkelundae sp. nov., which means that internal phragmocone features are still unknown in G. rosenkrantzi. Cyrtobelus hornbyense gen. nov, sp. nov. from the Campanian of western Canada constitutes the first record of early spirulids from the northeast Pacific, being based on seventeen extraordinarily well‐preserved phragmocones. This species differs from C. birkelundae sp. nov. only in the width of the siphuncular tube. The presence of a caecum, a nacre‐less conotheca that represents the continuation of the protoconch conotheca, conothecal flaps that anchor the mural parts of the septa, and a thin investment‐like sheath are characters shared only with Recent Spirula. In particular, the unusual protoconch architecture of Cyrtobelus gen. nov. challenges a phylogenetic origin within bactritoid‐like coleoids.     

Biological response to experimental damage of the phragmocone and siphuncle in Nautilus pompilius Linnaeus

Tsujino, Y & Shigeta, Y. 2012: Biological response to experimental damage of the phragmocone and siphuncle in Nautilus pompilius Linnaeus. Lethaia, Vol. 45, pp. 443–449. Three adult specimens of Nautilus pomplilius Linnaeus from the Philippines were experimented on to estimate the biological response to damage of the phragmocone and siphuncle in this cephalopod mollusc. In addition, the data obtained from the experiments were used for discussion of shell damage in ammonoids and in other extinct cephalopods. Specimen’s phragmocone and siphuncle were perforated and severed artificially, followed by observations in the laboratory tank during periods of 75 and 132 days. For at least 2 or 3 months, all individuals survived after damage to the phragmocone and siphuncle despite loss of neutral buoyancy. Based on our observations after completion of the experiments, the severed adoral remaining part of siphuncle healed by the siphunclar epithelium. In addition, perforation of the phragmocone was partly repaired by shell secretion from the dorsally extending mantle due to subsequent volution of shell growth. Our experiments revealed that damage to the phragmocone and siphuncle in Nautilus was not necessarily a lethal injury. It may be possible that such biological response also applies to extinct ammonoids and nautiloids. In a similar case of extinct ammonoids and nautiloids, damage to their phragmocone and siphuncle may also not have been a lethal injury as with Nautilus. However, some factors leading to death are likely to be dependent on the degree of damage to the phragmocone and siphuncle and influence of hydraulic pressure. □Ammonoids, injury, nautiloids, Nautilus, phragmocone, repair, siphuncle.     

Beak from the body chamber of an Early Carboniferous shelled longiconic coleoid cephalopod from Arkansas,USA

Here we report the discovery of an Early Carboniferous (Late Visean) 3D cephalopod beak displaying significant similarity to the lower beak of Recent coleoids. It was uncovered in a fragmentarily preserved, longiconic shell from the Moorefield Formation in Arkansas, USA. This shell comprises a fractured 29‐mm‐long body chamber having a maximum diameter of ~14 mm and showing an indistinct pro‐ostracum‐like structure. The beak‐bearing shell could easily have been mistaken for a bactritid or orthocerid if it were not for a coleoid‐type, weakly mineralized, evidently organic‐rich shell wall which shows a lamello‐columnar ultrastructure of a bulk of shell wall thickness and plate ultrastructure of thin outer layer. The specimen is assigned to an as‐yet unnamed shelled coleoid of a so far unknown high‐level taxonomic group. A partially exposed, 4.0‐mm‐long portion of the beak is the lower beak in oblique view from its left side. It exhibits fractured anthracite‐like black, apparently originally chitin material, helmet‐like general shape, broad hood with narrow shallow median groove and small notch posteriorly, pronounced pointed, non‐biomineralized upside belt rostrum, high shoulder and about a 90–100 degrees jaw angle. A broad hood and massive rostrum emphasize its similarity to the lower mandible of Recent Vampyroteuthis and signify that its unique, among living coleoids, structure has been existed for at least since Late Visean time (~333 my).     

The Evolution and Development of Cephalopod Chambers and Their Shape

The Ammonoidea is a group of extinct cephalopods ideal to study evolution through deep time. The evolution of the planispiral shell and complexly folded septa in ammonoids has been thought to have increased the functional surface area of the chambers permitting enhanced metabolic functions such as: chamber emptying, rate of mineralization and increased growth rates throughout ontogeny. Using nano-computed tomography and synchrotron radiation based micro-computed tomography, we present the first study of ontogenetic changes in surface area to volume ratios in the phragmocone chambers of several phylogenetically distant ammonoids and extant cephalopods. Contrary to the initial hypothesis, ammonoids do not possess a persistently high relative chamber surface area. Instead, the functional surface area of the chambers is higher in earliest ontogeny when compared to Spirula spirula. The higher the functional surface area the quicker the potential emptying rate of the chamber; quicker chamber emptying rates would theoretically permit faster growth. This is supported by the persistently higher siphuncular surface area to chamber volume ratio we collected for the ammonite Amauroceras sp. compared to either S. spirula or nautilids. We demonstrate that the curvature of the surface of the chamber increases with greater septal complexity increasing the potential refilling rates. We further show a unique relationship between ammonoid chamber shape and size that does not exist in S. spirula or nautilids. This view of chamber function also has implications for the evolution of the internal shell of coleoids, relating this event to the decoupling of soft-body growth and shell growth.     

Recent advances in heteromorph ammonoid palaeobiology

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.     

The locomotion system of Mesozoic Coleoidea (Cephalopoda) and its phylogenetic significance

A morphological comparison of shell‐muscle contacts in coleoid cephalopods mainly from the Early Jurassic (Toarcian) Posidonia Shales of Holzmaden (Germany), the Middle Jurassic (Callovian) Oxford Clay of Christian Malford (UK), Late Jurassic (Kimmeridgian‐Tithonian) plattenkalks of Solnhofen (Germany), and the Late Cretaceous (Cenomanian) of Hâdjoula and Hâkel (Lebanon) provides new and meaningful insights into their locomotion systems. The study shows that both pro‐ostracum‐ and gladius‐bearing coleoids are typified by a marginal mantle attachment and by distinctly separated fins, which usually insert (indirectly via the shell sac and basal fin cartilages) to posterior shell parts. While absent in gladius‐bearing forms, mantle‐locking cartilages might have existed already in pro‐ostracum‐bearing belemnoids. Similar to ectocochleate ancestors, funnel‐ and cephalic retractors are generally attached to the internal (ventral) shell surface. A comparison of Mesozoic and Recent gladius‐bearing coleoids shows that the locomotion system (most significantly the dorsal mantle configuration, and the presence of nuchal‐ and funnel‐locking cartilages) is fundamentally different. This does not support the concept of ‘fossil teuthids’, but suggests, owing to similarities with Recent Vampyroteuthis, placement of Mesozoic gladius‐bearing coleoids within the Octobrachia (Octopoda + Vampyromorpha). Classification of Mesozoic gladius‐bearing coleoids as octobrachians implies that: (1) unambiguous teuthids are still unknown in the fossil record and (2) the similarity between Recent and some fossil gladiuses represents a matter of homoplasy.     

Mosasaur predation on a nautiloid from the Maastrichtian Pierre Shale,Central Colorado,Western Interior Basin,United States

Mosasaurs were common predators on the ammonites that inhabited the upper water column of the Late Cretaceous Western Interior epicontinental seaway of North America. Mosasaurs developed predictable behaviour patterns for feeding on ammonite prey. There are no previous reports of mosasaur predation on the much less common Cretaceous nautiloids, possibly because of the prey's predominantly deep, epibenthic habitat, as deduced from modern Nautilus life habits. A single specimen of the highly inflated nautiloid, Eutrephoceras dekayi (Conrad), prey to a small adult mosasaur, likely Platycarpus, Prognathodon or Mosasaurus, is reported herein from the Pierre Shale of Colorado in the Early Maastrichtian biozone of Baculitesgrandis transitional to the biozone of B. clinolobatus. The nautiloid was killed in the same manner as described previously for discoid ammonites (Placenticeras, Sphenodiscus) from coeval strata in the USA and Canada.     

Post-mortem descent with septal implosion in Silurian nautiloids

Late Silurian nautiloids from Bohemia have either (1) thin and densely spaced septa of which many are broken and internally accumulated, or (2) thick and widely spaced septa wnich are all intact. Since the latest chamber (s) and the shell wall are undamaged, septal fragmentation occurred by implosion during postmortem sinking, with sea water rushing in through the siphuncle. The latest connecting ring(s) were more permeable than the immature ones, permitting pressure compensation in the latest chamber(s). Septal debris accumulated adapically indicating the (ultimate) sinking orientation. Depth (maxima) of the nautiloid habitats and of the Bohemian basin are estimated from the strength parameters of the broken and intact septa: brevicones — epipelagic, weak longicones — moderately shallow pelagic, strong longicones — nektobenthic, sea floor depth — several 100 m. Silurian-Nautiloids-Shell-Connecting rings-Bathymetry.     

The siphuncle in Georginidae and other Ordovician actinoceroid cephalopods

The lower Middle Ordovician carbonate sediments of the Georgina Basin in Australia contain many and varied Actinoceratida, including an endemic family, Georginidae Wade (1977), with siphuncular calcification consisting of radial lamellae separated by spaces (now sediment-filled) enclosed within calcified annulus walls. In each segment a distinct series of more massive calcareous engrafts grows inward from the inside of the connecting ring and is engrafted into adjacent annuli across the interannulus; this divides the perispatium into longitudinal perispatial sinuses. At each extremity each perispatial sinus is connected with passages leading through the segments to the axial space and, nearer to, or at the interannulus, with radial canals. Axial canals are few, but more than one is normal. Good preservation of Armenoceras and Actinoceras allows recognition of similar structures in the annuli of normal Actinoceratida.     

The structure of the chambered nautilus siphuncle: The siphuncular epithelium

The siphuncle of the chambered nautilus (Nautilus macromphalus) is composed of a layer of columnar epithelial cells resting on a vascularized connective tissue base. The siphuncular epithelium taken from chambers that have not yet begun to be emptied of cameral liquid has a dense apical brush border. The great number of apical cell junctions (zonula adherens) compared to the number of nuclei suggests extensive interdigitation of these cells. The perinuclear cytoplasm of these preemptying cells is rich in rough endoplasmic reticulum. The siphuncular epithelium of both emptying and “old” siphuncle (which has already completed emptying its chamber) both show little rough endoplasmic reticulum but do contain extensive systems of mitochondria-lined infoldings of the basolateral plasma membranes. Active transport of NaCl into the extracellular space of this tubular system probably entrains the water transport involved in the chamber-emptying process. Both emptying and old siphuncular epithelium also show large basal infoldings (canaliculi) continuous with the hemocoel, which appear to be filled with hemocyanin. The apical cell junctions of emptying and old siphuncular epithelium contain septate desmosomes that may help to prevent back-flow of cameral liquid into the chambers.     

Oxygen Isotope Variability within Nautilus Shell Growth Bands

Nautilus is often used as an analogue for the ecology and behavior of extinct externally shelled cephalopods. Nautilus shell grows quickly, has internal growth banding, and is widely believed to precipitate aragonite in oxygen isotope equilibrium with seawater. Pieces of shell from a wild-caught Nautilus macromphalus from New Caledonia and from a Nautilus belauensis reared in an aquarium were cast in epoxy, polished, and then imaged. Growth bands were visible in the outer prismatic layer of both shells. The thicknesses of the bands are consistent with previously reported daily growth rates measured in aquarium reared individuals. In situ analysis of oxygen isotope ratios using secondary ion mass spectrometry (SIMS) with 10 μm beam-spot size reveals inter- and intra-band δ¹⁸O variation. In the wild-caught sample, a traverse crosscutting 45 growth bands yielded δ¹⁸O values ranging 2.5‰, from +0.9 to -1.6 ‰ (VPDB), a range that is larger than that observed in many serial sampling of entire shells by conventional methods. The maximum range within a single band (~32 μm) was 1.5‰, and 27 out of 41 bands had a range larger than instrumental precision (±2 SD = 0.6‰). The results from the wild individual suggest depth migration is recorded by the shell, but are not consistent with a simple sinusoidal, diurnal depth change pattern. To create the observed range of δ¹⁸O, however, this Nautilus must have traversed a temperature gradient of at least ~12°C, corresponding to approximately 400 m depth change. Isotopic variation was also measured in the aquarium-reared sample, but the pattern within and between bands likely reflects evaporative enrichment arising from a weekly cycle of refill and replacement of the aquarium water. Overall, this work suggests that depth migration behavior in ancient nektonic mollusks could be elucidated by SIMS analysis across individual growth bands.     

Adhesive mechanisms in cephalopods: a review

Abstract

Several genera of cephalopods (Nautilus, Sepia, Euprymna and Idiosepius) produce adhesive secretions, which are used for attachment to the substratum, for mating and to capture prey. These adhesive structures are located in different parts of the body, viz. in the digital tentacles (Nautilus), in the ventral surface of the mantle and fourth arm pair (Sepia), in the dorsal epidermis (Euprymna), or in the dorsal mantle side and partly on the fins (Idiosepius). Adhesion in Sepia is induced by suction of dermal structures on the mantle, while for Nautilus, Euprymna and Idiosepius adhesion is probably achieved by chemical substances. Histochemical studies indicate that in Nautilus and Idiosepius secretory cells that appear to be involved in adhesion stain for carbohydrates and protein, whilst in Euprymna only carbohydrates are detectable. De-adhesion is either achieved by muscle contraction of the tentacles and mantle (Nautilus and Sepia) or by secretion of substances (Euprymna). The de-adhesive mechanism used by Idiosepius remains unknown.