Rhyncholites and conchorhynchs as calcified jaw elements in some late Cretaceous ammonites

Conspicuous calcareous coverings are present in the anterior region of 17 fossil jaws from late Cretaceous rocks of Hokkaido (Japan) and Sakhalin (U.S.S.R.). The jaws were preserved in calcareous nodules either in situ in body chambers of ammonites or in close association with identifiable ammonite conch remains. From the morphologic similarity between in situ and isolated jaws, they may be attributed to Tetragonites glabrus, Gaudryceras tenuiliratum, G. denseplicatum, G. sp., and Neophylloceras subramosum. The jaw apparatus of these species is composed of two three-dimensional black walls of carbonate apatite, which might be a diagenetic replacement of chitinous material. The calcareous coverings in both upper and lower jaws closely resemble those of upper (rhyncholite) and lower (conchorhynch) jaws of modern Nautilus as well as rhyncholite and conchorhynch fossils in their gross morphology, microstructure, and chemical composition. Calcified remains of cephalopod jaws known as rhyncholites and conchorhynchs have been reported from late Paleozoic to Recent. The present discovery of ammonoid rhyncholites and conchorhynchs suggests that at least some previously known late Paleozoic and Mesozoic counterparts belong to the Ammonoidea. The essential similarity of jaw elements of some Late Cretaceous ammonites and modern Nautilus gives reliable information on the feeding habits of the former. The sharp and thick ammonoid rhyncholites and conchorhynchs may have had a special function for cutting up food, similar to those of Nautilus.     

Cephalopod dynamic camouflage: bridging the continuum between background matching and disruptive coloration

Individual cuttlefish, octopus and squid have the versatile capability to use body patterns for background matching and disruptive coloration. We define—qualitatively and quantitatively—the chief characteristics of the three major body pattern types used for camouflage by cephalopods: uniform and mottle patterns for background matching, and disruptive patterns that primarily enhance disruptiveness but aid background matching as well. There is great variation within each of the three body pattern types, but by defining their chief characteristics we lay the groundwork to test camouflage concepts by correlating background statistics with those of the body pattern. We describe at least three ways in which background matching can be achieved in cephalopods. Disruptive patterns in cuttlefish possess all four of the basic components of ‘disruptiveness’, supporting Cott''s hypotheses, and we provide field examples of disruptive coloration in which the body pattern contrast exceeds that of the immediate surrounds. Based upon laboratory testing as well as thousands of images of camouflaged cephalopods in the field (a sample is provided on a web archive), we note that size, contrast and edges of background objects are key visual cues that guide cephalopod camouflage patterning. Mottle and disruptive patterns are frequently mixed, suggesting that background matching and disruptive mechanisms are often used in the same pattern.     

Why the ammonites snuffed it

Ammonites survived for millions of years despite steadily increasing competition from fish and coleoid cephalopods. The physiology and behaviour of Nautilus, the only remaining, if rather distantly related, ectocochleate cephalopod suggest as possible reasons the ability to remain aerobically active, albeit intermittently, at very low oxygen tensions and the ability to migrate vertically in and out of such zones at low cost. With the progressive oxygenation of the oceans shallow water hypoxic environments largely disappeared, trapping the ammonites and their vulnerable planktonic young stages between the depth limits imposed by their buoyancy mechanism and the high oxygen tension environments where they were exposed to faster and more economical predators and competitors.     

Mode of life and soft body shape of heteromorph ammonites

Ebel, K. 1992 04 15: Mode of life and soft body shape of heteromorph ammonites. Lethaia , Vol. 25, pp. 179–193. Oslo. ISSN 0024–1164.
Using the idea of a benthic mode of life for ammonites, based on a gastropod-like shell position, it is possible to reconstruct the development of all heteromorph ammonites by regarding single growth stages and the presumable acting forces. The reconstruction of shell formation, particularly the final shell position of heteromorphs with a hook, indicates that the soft body of the ammonite animal was considerably larger than comparison with the present-day Nautilus would suggest.     

Adaptable night camouflage by cuttlefish

Cephalopods are well known for their diverse, quick-changing camouflage in a wide range of shallow habitats worldwide. However, there is no documentation that cephalopods use their diverse camouflage repertoire at night. We used a remotely operated vehicle equipped with a video camera and a red light to conduct 16 transects on the communal spawning grounds of the giant Australian cuttlefish Sepia apama situated on a temperate rock reef in southern Australia. Cuttlefish ceased sexual signaling and reproductive behavior at dusk and then settled to the bottom and quickly adapted their body patterns to produce camouflage that was tailored to different backgrounds. During the day, only 3% of cuttlefish were camouflaged on the spawning ground, but at night 86% (71 of 83 cuttlefish) were camouflaged in variations of three body pattern types: uniform (n=5), mottled (n=33), or disruptive (n=34) coloration. The implication is that nocturnal visual predators provide the selective pressure for rapid, changeable camouflage patterning tuned to different visual backgrounds at night.     

Nautilus—a poor model for the function and behavior of ammonoids?

Inferences drawn from the biology, function, and behavior of closely related living forms facilitate interpretation of the mode of life of groups known only from the fossil record. The choice of phylogenetically relevant modern 'model organisms' can have critical bearing on the resulting interpretations. The biology and behavior of fossil ammonoids are often interpreted in the light of evidence derived from the study of modern Nautilus . However, examination of the fossil record and cladistic analyses both indicate that coleoids are much more closely related to ammonoids than is Nautilus . Coleoid biology and behavior differ dramatically from the biology and behavior of Nautilus . Thus, the inclusion of coleoids as examples, rather than reliance on Nautilus alone, produces a strikingly different vision of ammonoid biology and suggests that inferences of ammonoid biology and behavior that rely exclusively on Nautilus should be reviewed. Two features related to swimming ability in Nautilus , static stability and large retractor muscles, are much reduced in many ammonoids, leading to the interpretation that ammonoids were poorer swimmers than Nautilus . However, reexamination of the evidence indicates that static stability should not play a role in the swimming of ammonoids with long body chambers. In addition, functional arguments suggest that a coleoid-like swimming mechanism should have evolved prior to the loss of the body chamber in coleoids. Thus, a coleoid-like swimming mechanism is likely to have evolved prior to the separation of ammonoid and coleoid lineages. A mechanism is proposed by which a coleoid swimming mechanism, independent of retractor muscle size, could function in ammonoids with long body chambers.□ Ammonoids, ammonites, evolution, functional morphology , Nautilus, phylogeny .     

Beyond “living fossils”: Can comparative genomics finally reveal novelty?

Cephalopods have recently moved into the research focus due to the growing number of sequenced genomes, molecular tools, and laboratory culture (Albertin & Simakov, 2020)?. Genome data now allows us to ask how the many known novelties of cephalopod morphology are reflected in their genomes and gene regulation. A crucial gap in this understanding has been the limited information for the Nautilus, the last survivor of a cephalopod lineage that diverged from the highly derived coleoid clade (octopus, squid, cuttlefish) around 400 million years ago. The publication of Nautilus genomes (in this issue of Molecular Ecology [Huang et al., 2021; Zhang et al., 2021])? will help us understand which genetic changes happened when, and ultimately how they contributed to cephalopod evolution.     

Size and function of ammonite aptychi in comparison with buccal masses of modern cephalopods

Morton, Nicol & Nixon, Marion 1987 07 15: Size and function of ammonite aptychi in comparison with buccal masses of modem cephalopods.
Previous impressions that the size of ammonite aptychi is unusually large, thereby posing a problem to their interpretation as part of the jaw apparatus, are shown to be incorrect. The relative length of the aptychus for at least the Jurassic ammonite groups studied, at approximately 15% of the length of the body chamber, is remarkably constant in different taxonomic groups and sizes. This is well within the range for buccal mass length as percentage of mantle length of living cephalopods, being most similar to Octopus and Sepia and much smaller than Nautilus. The height and width of aptychi relative to whorl height and width are larger but again ammonites are probably not significantly different from modern cephalopods. The size and design, with no obvious structures for biting or crushing, suggest that ammonites were adapted to a particular type of relatively unspecialised feeding in which mostly small animals were ingested, possibly with some external digestion. The large shovel-like lower jaw may have functioned like a scoop for collecting large quantities of water and small prey, and movement of the buccal complex with the upper jaw almost closed against the lower jaw could have expelled water while retaining captured prey. Calcification of aptychi may have been protective, but more likely acted to weight the buccal mass for nektobenthic feeding and to make it more rigid.     

Flexible Spatial Orientation and Navigational Strategies in Chambered Nautilus

Nautilus is a remnant of an externally shelled cephalopod lineage that flourished between 450 and 60 million years ago. It is a deep‐water scavenger that lacks the complex brain and behavioural repertoire of its soft‐bodied relatives, the coleoid cephalopods. Nautilus makes repeated, nightly migrations from deep to shallow water along coral reef slopes to forage, thus an ability to navigate to known locations may be selectively advantageous. Alternatively, drifting passively with the current may be sufficient to locate food and mates distributed randomly over a large area. The derived neural structures that support learning and memory in coleoids are absent from the nautilus brain, indicating that substantial differences between learning abilities in nautilus and coleoids are likely. However, our previous work has demonstrated both associative and spatial learning in nautilus. In laboratory experiments, we tested whether Nautilus pompilius could learn to navigate towards a goal location using either visual cues or motor responses. Results indicate that animals relied both on proximate and distant visual cues to orient but did not use egocentric cues, a somewhat surprising finding given that nautilus spends most of its time in near darkness. Animals learned visual information rapidly and in some cases were able to switch to other tactics when salient visual cues were removed or manipulated.     

Communication and camouflage with the same 'bright' colours in reef fishes

Reef fishes present the observer with the most diverse and stunning assemblage of animal colours anywhere on earth. The functions of some of these colours and their combinations are examined using new non-subjective spectrophotometric measurements of the colours of fishes and their habitat. Conclusions reached are as follows: (i) the spectra of colours in high spatial frequency patterns are often well designed to be very conspicuous to a colour vision system at close range but well camouflaged at a distance; (ii) blue and yellow, the most frequently used colours in reef fishes, may be good for camouflage or communication depending on the background they are viewed against; and (iii) reef fishes use a combination of colour and behaviour to regulate their conspicuousness and crypsis.     

Über Nahrung und Ernährungsweise von Ammoniten

Until only a few years ago, the majority of ammonites were considered to have been rather swift nectonic swimmers of little ecologic bondage. About their food, nothing was known with certainty. Comparison with recent Coleoids and withNautilus suggests, however, that most ammonoids were poor swimmers. It is not even certain that the hyponome was used for locomotion as much as inNautilus. Slow locomotion made it impossible for ammonites to hunt for fish or other highly agile prey. Their lower jaws, which are identical to the structures known as aptychi and anaptychi (formerly considered to be opercula), are like shovels and make it impossible for their bearers actually to bite and cut with them. Consequently the structure of their jaws also prevented their hunting large and active prey. Their jaws were probably actually used like shovels along the bottom of the sea. They may have gathered small or slow benthonic organisms like foraminifers, ostracodes, small crustaceans, young brachiopods, corals, bryozoa, but also drifting or slowly swimming and dead organisms. Several unusually well preserved stomachs of ammonites contained foraminifers, ostracodes, crinoids, smaller ammonites, and possibly inarticulate brachiopods. It is concluded that, ecologically, ammonites inhabited the place of modern bottom-feeding gastropods.     

To be seen or to hide: visual characteristics of body patterns for camouflage and communication in the Australian giant cuttlefish Sepia apama

It might seem obvious that a camouflaged animal must generally match its background whereas to be conspicuous an organism must differ from the background. However, the image parameters (or statistics) that evaluate the conspicuousness of patterns and textures are seldom well defined, and animal coloration patterns are rarely compared quantitatively with their respective backgrounds. Here we examine this issue in the Australian giant cuttlefish Sepia apama. We confine our analysis to the best-known and simplest image statistic, the correlation in intensity between neighboring pixels. Sepia apama can rapidly change their body patterns from assumed conspicuous signaling to assumed camouflage, thus providing an excellent and unique opportunity to investigate how such patterns differ in a single visual habitat. We describe the intensity variance and spatial frequency power spectra of these differing body patterns and compare these patterns with the backgrounds against which they are viewed. The measured image statistics of camouflaged animals closely resemble their backgrounds, while signaling animals differ significantly from their backgrounds. Our findings may provide the basis for a set of general rules for crypsis and signals. Furthermore, our methods may be widely applicable to the quantitative study of animal coloration.     

Vertical distribution and migration patterns of Nautilus pompilius

Vertical depth migrations into shallower waters at night by the chambered cephalopod Nautilus were first hypothesized early in the early 20(th) Century. Subsequent studies have supported the hypothesis that Nautilus spend daytime hours at depth and only ascend to around 200 m at night. Here we challenge this idea of a universal Nautilus behavior. Ultrasonic telemetry techniques were employed to track eleven specimens of Nautilus pompilius for variable times ranging from one to 78 days at Osprey Reef, Coral Sea, Australia. To supplement these observations, six remotely operated vehicle (ROV) dives were conducted at the same location to provide 29 hours of observations from 100 to 800 meter depths which sighted an additional 48 individuals, including five juveniles, all deeper than 489 m. The resulting data suggest virtually continuous, nightly movement between depths of 130 to 700 m, with daytime behavior split between either virtual stasis in the relatively shallow 160-225 m depths or active foraging in depths between 489 to 700 m. The findings also extend the known habitable depth range of Nautilus to 700 m, demonstrate juvenile distribution within the same habitat as adults and document daytime feeding behavior. These data support a hypothesis that, contrary to previously observed diurnal patterns of shallower at night than day, more complex vertical movement patterns may exist in at least this, and perhaps all other Nautilus populations. These are most likely dictated by optimal feeding substrate, avoidance of daytime visual predators, requirements for resting periods at 200 m to regain neutral buoyancy, upper temperature limits of around 25°C and implosion depths of 800 m. The slope, terrain and biological community of the various geographically separated Nautilus populations may provide different permutations and combinations of the above factors resulting in preferred vertical movement strategies most suited for each population.     

Thriving at high hydrostatic pressure: the example of ammonoids (extinct cephalopods)

Ammonoids are a group of extinct mollusks belonging to the same class of the living genus Nautilus (cephalopoda). In both Nautili and ammonoids, the (usually planospiral) shell is divided into chambers separated by septa that, during their lifetime, are filled with gas at atmospheric pressure. The intersection of septa with the external shell generates a curve called the suture line, which in living and most fossil Nautili is fairly uncomplicated. In contrast, suture lines of ancient ammonoids were gently curved and during the evolution of the group became highly complex, in some cases so extensively frilled as to be considered as fractal curves. Numerous theories have been put forward to explain the complexity of suture ammonoid lines. Calculations presented here lend support to the hypothesis that complex suture lines aided in counteracting the effect of the external water pressure. Additionally, it is suggested that complex suture lines diminished shell shrinkage caused by water pressure, and thus aided in improving buoyancy. Understanding the reason for complex sutures in ammonoids represents an important issue in paleobiology with potential applications to the problem of the resistance of hollow mechanical structures subjected to high pressure.     

A fish‐eye view of cuttlefish camouflage using in situ spectrometry

Cuttlefish are colour blind yet they appear to produce colour‐coordinated patterns for camouflage. Under natural in situ lighting conditions in southern Australia, we took point‐by‐point spectrometry measurements of camouflaged cuttlefish, Sepia apama, and various natural objects in the immediate visual surrounds to quantify the degree of chromatic resemblance between cuttlefish and backgrounds to potential fish predators. Luminance contrast was also calculated to determine the effectiveness of cuttlefish camouflage to this information channel both for animals with or without colour vision. Uniform body patterns on a homogeneous background of algae showed close resemblance in colour and luminance; a Uniform pattern on a partially heterogeneous background showed mixed levels of resemblance to certain background features. A Mottle pattern with some disruptive components on a heterogeneous background showed general background resemblance to some benthic objects nearest the cuttlefish. A noteworthy observation for a Disruptive body pattern on a heterogeneous background was the wide range in spectral contrasts compared to Uniform and Mottle patterns. This suggests a shift in camouflage tactic from background resemblance (which hinders detection by the predator) to more specific object resemblance and disruptive camouflage (which retards recognition). © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 535–551.     

The occurrence and preservation of ammonites in the Blue Lias Formation (lower Jurassic) of Devon and Dorset, England and their palaeoecological, sedimentological and diagenetic significance

More than two thirds of beds in the lowest Jurassic, Blue Lias Formation lack ammonites, which are commonly preserved in irregular or planar-bedded, bioturbated limestones, very rarely in laminated limestones and almost never in laminated black shales. Ammonites are preserved in 3D in nodular and planar-bedded limestones and at any orientation to bedding. Co-occurrence with macrobenthos and absence from beds without benthos suggest that Blue Lias ammonites were nektobenthonic. Scour structures and imbrication of ammonites in the Best Bed imply presence of traction currents. Lack of epifauna on large cephalopod shells (and other fossils) implies rapid deposition in event beds. Blue Lias deposition was episodic, not slow and continuous as the fine grain size implies. Undistorted trace fossils, uncrushed ammonites and stable isotope values all suggest early cementation of limestone beds from pore waters of a similar composition to contemporary Jurassic sea water. A clear diagenetic trend exists, with limestones having least, and laminated black shales most, modified stable isotope values. Contrast between trace fossil fills and host sediment demonstrates that Blue Lias rhythms are primary, but limestone beds have been diagenetically cemented.     

Ecological,behavioral, and phylogenetic influences on the evolution of dorsal color pattern in geckos*

The dorsal surfaces of many taxonomic groups often feature repetitive pattern elements consisting of stripes, spots, or bands. Here, we investigate how distinct categories of camouflage pattern work by relating them to ecological and behavioral traits in 439 species of gecko. We use phylogenetic comparative methods to test outstanding hypotheses based on camouflage theory and research in other taxa. We found that bands are associated with nocturnal activity, suggesting bands provide effective camouflage for motionless geckos resting in refugia during the day. A predicted association between stripes and diurnal activity was not supported, suggesting that stripes do not work via dazzle camouflage mechanisms in geckos. This, along with a lack of support for our prediction that plain patterning should be associated with open habitats, suggests that similar camouflage patterns do not work in consistent ways across taxa. We also found that plain and striped lineages frequently switched between using open or closed habitats, whereas spotted lineages rarely transitioned. This suggests that pattern categories differ in how specialized or generalized their camouflage is. This result has ramifications for theory on how camouflage compromises to background heterogeneity and how camouflage pattern might influence evolutionary trajectories.     

Animal behaviour and algal camouflage jointly structure predation and selection

Trait variation can structure interactions between individuals, thus shaping selection. Although antipredator strategies are an important component of many aquatic systems, how multiple antipredator traits interact to influence consumption and selection remains contentious. Here, I use a common larval dragonfly (Epitheca canis) and its predator (Anax junius) to test for the joint effects of activity rate and algal camouflage on predation and survival selection. I found that active and poorly camouflaged Epitheca were more likely to be consumed, and thus, survival selection favoured inactive and well‐camouflaged individuals. Notably, camouflage dampened selection on activity rate, likely by reducing attack rates when Epitheca encountered a predator. Correlational selection is therefore conferred by the ecological interaction of traits, rather than by opposing selection acting on linked traits. I suggest that antipredator traits with different adaptive functions can jointly structure patterns of consumption and selection.     

Mollusk shell formation: mapping the distribution of organic matrix components underlying a single aragonitic tablet in nacre

Control over mineral formation in mollusk shells is exerted by the macromolecules of the organic matrix. Using histochemical methods, we mapped the carboxylates and sulfates of proteins and polysaccharides on the surfaces of decalcified interlamellar matrices from the nacreous shell layer of the cephalopod Nautilus pompilius, expanding upon an earlier study by Crenshaw and Ristedt [Crenshaw, M.A., Ristedt, H., 1976. The histochemical localization of reactive groups in septal nacre from Nautilus pompilius. In: Watabe, N., Wilbur, K.M. (Ed.), The Mechanisms of Mineralization in the Invertebrates and Plants. University of South Carolina Press, Colombia, pp. 355-367]. We observed four different zones underlying a single crystal: (1) a central spot rich in carboxylates; (2) a central ring-shaped area rich in sulfates; (3) an area between the central nucleation region and the imprint periphery containing carboxylates, and (4) the intertabular matrix, rich in carboxylates and sulfates. We also mapped matrix functional groups on the nacreous matrix surfaces of the bivalve Atrina rigida, but did not identify well-defined zones. Immuno-mapping of the constituents of the aragonite-nucleating protein fraction from Atrina nacre showed that these macromolecules are located both in the intertabular matrix and in the center of the crystal imprints for both Atrina and Nautilus matrix surfaces. Their presence at the latter location is consistent with their purported role in aragonite nucleation. The observed differentiation in the distribution of matrix components and their functional groups shows that the different stages of single crystal growth are highly controlled by the matrix.