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 .     

Buccal mass structure of the Cretaceous ammonite Gaudryceras

A rhynchaptychus attributed to the lower jaw of Gaudryceras sp. from the Upper Santonian rocks of Hokkaido, Japan, has an interesting impression well-preserved on the inner surface of the outer lamella. The impression is regarded as the imprints of chitin-secreting cells (beccublast cells), because of similarity in the characteristic arrangement of polygonal pits to those of modern coleoids. Each unit cell impression of G. sp. is, however, about five to ten times larger than in modern coleoids known to us. In modern coleoids and Nautilus a layer of tall beccublast cells is intercalated between the buecal muscles and the outer side of the upper jaw and/or the inner side of the lower jaw. The other sides of the jaws are, in contrast, free from jaw muscles, and are covered directly with a thin connecting tissue. Based on these observations a possible buecal mass structure of G. sp. is restored. The beccublast cell impressions of Gaudryceras and modern coleoids markedly differ from that of modern Nautilus in the absence of numerous micropores. This fact suggests weaker mechanical properties of the jaw muscles in Gaudryceras than in Nautilus , as the branching ends of beccublast cells of the latter are inserted in the micropores to keep a firm attachment of the jaw muscles on the jaw plates.     

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.     

Kompass im Kopf

Ant compass – how desert ants learn to navigate Successful spatial orientation is a daily challenge for many animals. Cataglyphis desert ants are famous for their navigational performances. They return to the nest after extensive foraging trips without any problems. How do ants take their navigational systems into operation? After conducting different tasks in the dark nest for several weeks, they become foragers under bright sun light. This transition requires both a drastic switch in behavior and neuronal changes in the brain. Experienced foragers mainly rely on visual cues. They use a celestial compass and landmark panoramas. For that reason, naïve ants perform stereotype learning walks to calibrate their compass systems and acquire information about the nest's surroundings. During their learning walks, the ants frequently look back to the nest entrance to learn the homing direction. For aligning their gazes, they use the earth's magnetic field as a compass reference. This magnetic compass in Cataglyphis ants was previously unknown.     

A revisited phylogeography of Nautilus pompilius

The cephalopod genus Nautilus is considered a “living fossil” with a contested number of extant and extinct species, and a benthic lifestyle that limits movement of animals between isolated seamounts and landmasses in the Indo‐Pacific. Nautiluses are fished for their shells, most heavily in the Philippines, and these fisheries have little monitoring or regulation. Here, we evaluate the hypothesis that multiple species of Nautilus (e.g., N. belauensis, N. repertus and N. stenomphalus) are in fact one species with a diverse phenotypic and geologic range. Using mitochondrial markers, we show that nautiluses from the Philippines, eastern Australia (Great Barrier Reef), Vanuatu, American Samoa, and Fiji fall into distinct geographical clades. For phylogenetic analysis of species complexes across the range of nautilus, we included sequences of Nautilus pompilius and other Nautilus species from GenBank from localities sampled in this study and others. We found that specimens from Western Australia cluster with samples from the Philippines, suggesting that interbreeding may be occurring between those locations, or that there is limited genetic drift due to large effective population sizes. Intriguingly, our data also show that nautilus identified in other studies as N. belauensis, N. stenomphalus, or N. repertus are likely N. pompilius displaying a diversity of morphological characters, suggesting that there is significant phenotypic plasticity within N. pompilius.     

Visual cues override olfactory cues in the host-finding process of the monophagous leaf beetle Altica engstroemi

It is generally assumed that specialist insect herbivores utilize plant odours to find their particular host plants and that visual cues are of minor importance in the host‐finding process. We performed Y‐tube olfactometer bioassays and small‐scale field experiments to determine whether, under laboratory and field conditions, the monophagous herbivore Altica engstroemi J. Sahlberg (Coleoptera: Chrysomelidae: Alticinae) is guided to its host plant Filipendula ulmaria (L.) Maxim. (Rosaceae) by visual or olfactory cues. The olfactometer tests showed that A. engstroemi was never attracted to odours, either from undamaged or from damaged plants. Even starvation for 24 h did not change this behaviour. However, the field experiment showed that visual cues alone were sufficient to attract a significant number of starved beetles when offered a choice between bagged host plants and bagged green plastic control ‘plants’. Our findings contrast with the general view that plant odours constitute the major cue in the host‐finding process among specialized phytophagous insects. A review of the literature for the period 1986–2006 inclusive, relating to host‐plant finding in Chrysomelidae, identified studies of 19 chrysomelid species, all of which were guided by olfactory cues. No species were guided to their host by visual cues. Although some studies demonstrated that chrysomelids may exhibit orientation responses to colour or contrast, our study on A. engstroemi is the only one demonstrating that visual cues affect host‐plant selection in a chrysomelid species. We suggest that the use of visual cues in host‐finding may evolve among chrysomelids with limited dispersal ability in persistent habitats and may be found among species monophagous on abundant host plants that dominate the structure of the plant community, that is, where the host plant's presence is predictable in time and space.     

Multisensory Control of Multimodal Behavior: Do the Legs Know What the Tongue Is Doing?

Understanding of adaptive behavior requires the precisely controlled presentation of multisensory stimuli combined with simultaneous measurement of multiple behavioral modalities. Hence, we developed a virtual reality apparatus that allows for simultaneous measurement of reward checking, a commonly used measure in associative learning paradigms, and navigational behavior, along with precisely controlled presentation of visual, auditory and reward stimuli. Rats performed a virtual spatial navigation task analogous to the Morris maze where only distal visual or auditory cues provided spatial information. Spatial navigation and reward checking maps showed experience-dependent learning and were in register for distal visual cues. However, they showed a dissociation, whereby distal auditory cues failed to support spatial navigation but did support spatially localized reward checking. These findings indicate that rats can navigate in virtual space with only distal visual cues, without significant vestibular or other sensory inputs. Furthermore, they reveal the simultaneous dissociation between two reward-driven behaviors.     

Relevance of visual and olfactory cues for host location in the mustard leaf beetle Phaedon cochleariae

The relevance of visual and olfactory cues for host‐plant location is investigated in males and females of the oligophagous mustard leaf beetle Phaedon cochleariae Fabricius (Coleoptera: Chrysomelidae). Different objects are offered in a walking arena and the behaviour of beetles is observed. Beetles orient toward vertically or horizontally striped black and white pattern independent of stripe orientation. The results suggest that contrast facilitates orientation in the field, whereas the pattern itself may be less important for host location in dense vegetation. The response to green and yellow objects is tested to investigate discrimination abilities between young (green) and mature (yellow) leaves. Beetles prefer green over yellow independent of material (cardboard or leaves of Nasturtium officinale R. Br., Brassicaceae). Preference behaviour tested in a dual‐choice contact assay coincides with visual preferences, where adults prefer young, more nutritious leaves for feeding and oviposition. Furthermore, females discriminate between visual cues of green leaves and green cardboard, whereas males do not, indicating that females are more sensitive in colour discrimination. Differences in colour wavelength influence the choice of beetle behaviour more strongly than differences in intensity. Both sexes of P. cochleariae prefer volatiles of the host plant N. officinale, whereas only females respond to the main volatile compound 2‐phenylethyl isothiocyanate. Given a choice between visual and olfactory cues, males orientate towards the colour cues, whereas females do not show any preferences. In males, visual cues may thus override olfactory cues, whereas, in females, both are equally important, which may reflect different ecological requirements and/or physiological abilities.     

Visual Discrimination of Kin in Mandrills

Phenotype matching, a learning mechanism that evolved based on phenotypic cues shared among relatives, may provide animals with the ability to recognize unfamiliar kin. The generalization of this mechanism across animal species is debated, however, because appropriate tests are difficult to design due to possible confounding effects of familiarity. Hence, only a few studies have examined evidence for the existence of such a mechanism in natural populations. Here, we tested the phenotype matching hypothesis based on visual cues in a semi‐free‐ranging population of mandrills (Mandrillus sphinx) that contains individuals related to different degrees and where familiarity is controlled for. Using an experimental design based on the presentation of photographs, we show that mandrills discriminate unfamiliar relatives using facial cues alone. Our results build on earlier studies, showing that primates use phenotype matching to recognize and subsequently discriminate unfamiliar kin. We suggest that facial features along with other visual and non‐visual cues provide a proximate mechanism for kin selection to operate.     

Influence of habitat complexity on route learning among different populations of climbing perch (Anabas testudineus Bloch, 1792)

Animals use different behavioral strategies to maximize their fitness in the natural environment. Learning and memory are critical in this context, allowing organisms to flexibly and rapidly respond to environmental changes. We studied how the physical characteristics of the native habitat influence the spatial learning capacity of Anabas testudineus belonging to four different populations collected from two streams and two ponds, in a linear maze. Stream fish were able to learn the route faster than pond fish irrespective of the presence or absence of landmarks in the maze. However, climbing perch collected from ponds learned the route faster in the maze provided with landmarks than in Plain maze. The results indicate that fish inhabiting a lotic ecosystem use egocentric cues in route learning rather than visual cues like landmarks. A local landmark may be a more reliable cue in route learning in a relatively stable habitat like a pond. In flowing aquatic systems, water flow may continually disrupt the visual landscape and thus landmarks as visual cues become unreliable. Spatial learning is thus a fine-tuned response to the complexity of the habitat and early rearing conditions may influence the spatial learning ability in fish.     

Role of chemical and visual cues in food recognition by leatherback posthatchlings (Dermochelys coriacea L)

We raised leatherback posthatchlings in the laboratory for up to 7 weeks to study the role of visual and chemical cues in food recognition and food-seeking behavior. Turtles were reared on a formulated (artificial gelatinous) diet and had no contact with test materials until experiments began. Subjects were presented with visual cues (a plastic jellyfish; white plastic shapes [circle, square, diamond] similar in surface area to the plastic model), chemical cues (homogenates of lion's mane jellyfish, Cyanea capillata; moon jellyfish, Aurelia aurita; and a ctenophore, Ocyropsis sp., introduced through a water filter outflow), and visual and chemical cues presented simultaneously. Visual stimuli evoked an increase in swimming activity, biting, diving, and orientation toward the object. Chemical cues elicited an increase in biting, and orientation into water currents (rheotaxis). When chemical and visual stimuli were combined, turtles ignored currents and oriented toward the visual stimuli. We conclude that both cues are used to search for, and locate, food but that visual cues may be of primary importance. We hypothesize that under natural conditions turtles locate food visually, then, as a consequence of feeding, associate chemical with visual cues. Chemical cues then may function alone as a feeding attractant.     

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.     

Reef waters stimulate substratum exploration in planulae from brooding Caribbean corals

This study tested the hypothesis that waters surrounding reefs with healthy coral populations are more likely than degraded sites to induce planulae to navigate downward and begin benthic probing. In the laboratory, larvae from two brooding Caribbean coral species, Agaricia tenuifolia and Porites astreoides, were introduced to seawater collected at (1) 1 m above shallow, healthy reef with high-coral cover, (2) 1 m above shallow, degraded reef with high-macroalgal cover, and (3) ~400 m ocean-ward of the reef in deep, blue water. Counter to the hypothesis, water from both the healthy and degraded reef caused the larvae to swim downward and begin benthic probing. These results suggest that substances carried in reef waters may contribute to macro-scale habitat selection by planulae and that understanding how these waterborne cues mesh with other stimuli used by planulae to select a settlement site may be valuable for deciphering a site’s recruitment potential for corals.     

Female assessment of male functional fertility during mate choice in a promiscuous fish

Sexual selection should favour females that can assess the functional fertility of available sexual partners and avoid mating with recently mated, sperm‐depleted males. Our current understanding of the sensory mechanism(s) underlying female assessment of males based on their functional fertility and avoidance of sperm‐depleted males is incomplete. Female Trinidadian guppies (Poecilia reticulata) are known to avoid mating with males that they had previously observed mating with other females. Here, we investigated experimentally the proximate sensory cues that they use to distinguish between paired size‐ and colour‐matched mated and unmated males in the absence of visual public information on their prior mating histories. When only water‐borne chemical cues from the males were available, females avoided the previously mated male and preferred the unmated one, but they chose randomly when only male visual cues (and no chemical cues) were available. They also preferred unmated over mated males when freely swimming with them in a more sensorially complex environment with multiple male cues (i.e., visual, chemical and mechanical cues) concurrently available. Females exhibited no preference for either stimulus males when both were unmated, irrespectively of the sensory environment. These novel results suggest that, in the absence of prior visual public information on the recent mating histories of males, female guppies use olfactory cues putatively emitted by mated males to avoid mating with them. The source and nature of the implicated olfactory cues and the fitness benefits gained by female guppies in sexually preferring males that have not recently mated remain unknown and warrant further research.     

Turbidity Hampers Mate Choice in a Pipefish

European coastal waters have in recent years become more turbid as algal growth has increased, probably due to eutrophication, global warming and changes in fish communities. Turbidity reduces visibility, and such changes may in turn affect animal behaviour as well as evolutionary processes that are dependent on visual stimuli. In this study we experimentally manipulated water visibility and olfactory cues to investigate mate choice using the sex role‐reversed broad‐nosed pipefish Syngnathus typhle as our study organism. We show that males spent significantly longer time assessing females when they had access to full visual cues, compared to when visibility was reduced. Presence or absence of olfactory cues from females did not affect mate choice, suggesting that the possible use of smell could not make up for a reduction in visibility. This implies that mate choice is environmentally dependent and that an increased turbidity may affect processes of sexual selection through an impaired possibility for visually based mate choice.     

Personality effects on spatial learning: Comparisons between visual conditions in a weakly electric fish

Recent research has explored links between cognition and personality, with prominent hypotheses proposing that personality drives consistent individual differences in cognitive function. These hypotheses particularly expect bolder individuals to be faster, but less accurate, as a trade‐off in cognitive function. However, cognitive processes are typically interconnected and defined in more complex terms than simply speed and accuracy. Here, we present evidence that personality‐based differences in learning rates are a result of differences in decision‐making during training in a two‐alternative forced‐choice spatial memory task. This was examined in the mormyrid fish Gnathonemus petersii in the presence of light, where both vision and the electric sense are available, and in the dark, where visibility is limited and fish rely mostly on electrosensing. The species is adapted for the dark to avoid visual predators; thus, the presence of light can induce high‐risk and the dark low‐risk. We show that light conditions had little effect on learning, with bolder fish learning faster both in the light and in the dark conditions. Yet the relationship between learning rates and error rates indicates that the effect on learning is indirectly influenced by accuracy during training. Speed‐accuracy trade‐offs were not found in decision‐making, with bolder individuals deciding faster and more accurately both in the light and in the dark. Only learning strategy was affected by light conditions, with significantly more fish preferring response to place learning in the dark than in the light, where distal cues were not visible. We conclude that other than effects from the integration of visual information, bolder individuals show a consistently greater tendency to explore and find food rewards during training. This affects their decision‐making and in turn their learning performance. We highlight the complexity by which personality‐based effects are exhibited in spatial associative learning.     

Plasticity between visual input pathways and the head direction system

Animals can maintain a stable sense of direction even when they navigate in novel environments, but how the animal's brain interprets and encodes unfamiliar sensory information in its navigation system to maintain a stable sense of direction is a mystery. Recent studies have suggested that distinct brain structures of mammals and insects have evolved to solve this common problem with strategies that share computational principles; specifically, a network structure called a ring attractor maintains the sense of direction. Initially, in a novel environment, the animal's sense of direction relies on self-motion cues. Over time, the mapping from visual inputs to head direction cells, responsible for the sense of direction, is established via experience-dependent plasticity. Yet the mechanisms that facilitate acquiring a world-centered sense of direction, how many environments can be stored in memory, and what visual features are selected, all remain unknown. Thanks to recent advances in large scale physiological recording, genetic tools, and theory, these mechanisms may soon be revealed.     

Navigation in Familiar Environments by the Weakly Electric Elephantnose Fish,Gnathonemus petersii L. (Mormyriformes,Teleostei)

Gnathonemus petersii use electrolocation to navigate in unfamiliar environments. The goal of these experiments was to determine whether fish could learn the location of a fixed aperture after interference with selected sensory input. By manipulating environmental cues (aperture height and water depth) and comparing the fish's performance, the contributions of the electrosensory system, vision, and hydrostatic pressure were examined. The fish's task was to find a circular aperture in a wall dividing a 200-litre aquarium into two equal compartments. In experiment 1, the position of the aperture was raised by 10.1 cm after the fish had become familiar with its original location. In experiment 2, the water level was raised by 10 cm (leaving the aperture unchanged). When the aperture was raised, intact fish found the new aperture with no difficulty, whereas blind, electrically 'silent', and sham-operated fish were slow finding the new position. When the water level was raised, all fish increased the height at which they contacted the wall, increased their electric-organ discharge (EOD) rate, and located the aperture. This increase, in response to the rapid change in water depth, suggests that all fish used hydrostatic pressure cues to maintain depth orientation, and that those fish that learned the aperture height had used hydrostatic cues to locate its position. The data suggest that G. petersii develop an internal representation based on an electrosensory central expectation and hydrostatic cues. The fish develop a sensory 'image' of their immediate environment and associate a specific image with a specific depth. As the environment becomes more familiar, the fish apparently attend less to electrosensory information and navigate according to the internal representation, relying primarily on hydrostatic pressure cues.     

A molecular and karyological approach to the taxonomy of Nautilus

Nautiloids, the externally shelled cephalopods of Cambrian origin, are the most ancient lineage among extant cephalopods. Their ancestral characters are explored based on morphological and molecular data (18S rDNA sequence) to investigate the evolution of present cephalopod lineages. Among molluscs, nautilus 18S rDNA gene is the longest reported so far, due to large nucleotidic insertions. By comparison with other 18S sequences, the complete gene of N. macromphalus helps to clarify the taxonomic status of the three universally recognised Nautilus species. The range of interspecific molecular differences supports separation of the present species into two surviving ectocochleate genera, Nautilus and Allonautilus. Nautiloid 18S is considered as corresponding to the ancestral form of 18S as is the number of chromosomes in Nautilus (52), the lowest among cephalopods. Comparison of karyological characteristics amongst cephalopods in a phylogenetic context suggests a possible correlation between duplication events and lineage divergence.     

Lack of species discrimination based on chemical cues by male sailfin mollies, <Emphasis Type="Italic">Poecilia latipinna</Emphasis>

When making mating decisions, individuals may rely on multiple cues from either the same or multiple sensory modalities. Although the use of visual cues in sexual selection is well studied, fewer studies have examined the role of chemical cues in mate choice. In addition, few studies have examined how visual and/or chemical cues affect male mating decisions. Male mate choice is important in systems where males must avoid mating with heterospecific females, as is found in a mating complex of Poecilia. Male sailfin mollies, Poecilia latipinna, are sexually parasitized by gynogenetic Amazon mollies, P. formosa. Little is known about the mechanism by which male sailfin mollies base their mating decisions. Here we tested the hypothesis that male sailfin mollies from an allopatric and a sympatric population with Amazon mollies use multiple cues to distinguish between conspecific and heterospecific females. We found that male sailfin mollies recognized the chemical cues of conspecific females, but we found no support for the hypothesis that chemical cues are by themselves sufficient for species discrimination. Lack of discrimination based on chemical cues alone may be due to the close evolutionary history between P. latipinna and P. formosa. Males from populations sympatric with Amazon mollies did not differentially associate with females of either of the two species when given access to both visual and chemical cues of the females, yet males from the allopatric population did associate more with conspecific females than with heterospecific females in the presence of both chemical and visual cues. The lack of discrimination by males from the sympatric population between conspecific and heterospecific females based on both chemical and visual cues suggests that these males require more complex combinations of cues to distinguish species, possibly due to the close relatedness of these species.