Sound production by the western rock lobster Panulirus longipes (Milne Edwards)

Sound characteristics and sound emitting structures of larval, juvenile and adult rock lobsters, Panulirus longipes (Milne Edwards) have been investigated. There is a linear correlation between carapace length and dimensions of the sound producing organ (number and size of plectrum ridges) as well as average duration of a single squeak. The number of pulses per 100 ms squeak, however, decreases asymptotically from $\text{\$}\text{?}$30 in larval forms to about 10 in mature adults. Behavioural changes accompany this process. The energy of a single pulse is distributed equally over the audio frequency range. The same is true for the ‘slow rattle’ sound, produced by mandible-grinding during feeding or moments of extreme agitation. Any predator attacking a rock lobster, is likely to receive injuries from the outstretched, spiny antennae. It is concluded that the antennal sound, accompanying the rock lobsters defence, functions as a warning device aimed primarily at predators, which will learn to associate the sound of rock lobsters with the unpleasant experience of the sharp antennal spines.     

Towards a Supertree of Arthropoda: A Species-Level Supertree of the Spiny,Slipper and Coral Lobsters (Decapoda: Achelata)

While supertrees have been built for many vertebrate groups (notably birds, mammals and dinosaurs), invertebrates have attracted relatively little attention. The paucity of supertrees of arthropods is particularly surprising given their economic and ecological importance, as well as their overwhelming contribution to biodiversity. The absence of comprehensive archives of machine-readable source trees, coupled with the need for software implementing repeatable protocols for managing them, has undoubtedly impeded progress. Here we present a supertree of Achelata (spiny, slipper and coral lobsters) as a proof of concept, constructed using new supertree specific software (the Supertree Toolkit; STK) and following a published protocol. We also introduce a new resource for archiving and managing published source trees. Our supertree of Achelata is synthesised from morphological and molecular source trees, and represents the most complete species-level tree of the group to date. Our findings are consistent with recent taxonomic treatments, confirming the validity of just two families: Palinuridae and Scyllaridae; Synaxidae were resolved within Palinuridae. Monophyletic Silentes and Stridentes lineages are recovered within Palinuridae, and all sub-families within Scyllaridae are found to be monophyletic with the exception of Ibacinae. We demonstrate the feasibility of building larger supertrees of arthropods, with the ultimate objective of building a complete species-level phylogeny for the entire phylum using a divide and conquer strategy.     

Morphology and distribution of antennular setae of scyllarid lobsters (Scyllarides aequinoctialis, S. latus, and S. nodifer) with comments on their possible function

Abstract. Lateral flagella of the antennules of scyllarid lobsters were examined for setal morphology and distribution via scanning electron microscopy. Setal distribution patterns were mapped directly for 3 regions of the antennule ( base, tuft , and tip ) and analyzed for differences: (1) between left and right antennules, (2) between males and females within a species, and (3) among species by comparing counts of setae per annulus in the ventral tuft region only. Six types of antennular setae were identified based on their external morphology: aesthetases, simple, modified simple, asymmetric, hemi-plumose, and toothbrush setae. These different types were organized in a clear pattern over the ventral and dorsal surfaces of the lateral flagella of the antennule. Aesthetase, asymmetric, modified simple, and hemi-plumose setae were found only on annuli in the tuft region between the distal and proximal ends of the flagellum. Simple setae were found on all annuli of all regions of the antennule, and toothbrush setae were mainly concentrated on all annuli of the base region and on proximal annuli of the tuft region . All species of scyllarids examined had the same general pattern of setal distribution and no differences were found between left and right, or male and female antennules. Similar setae located on the lateral antennules of species from the families Nephrophidae and Palinuridae (clawed and spiny lobsters) have been previously described as chemo- and/or mechanoreceptive for use in distance chemoreception (i.e., detection and orientation to olfactory stimuli). Based on work on clawed and spiny lobsters, we predict that the aesthetases on slipper lobsters have a chemoreceptive function and that simple and toothbrush setae may have a bimodal chemo- and mechanoreceptive function.     

An evaluation of lipid‐ and morphometric‐based indices of nutritional condition for early benthic stage spiny lobsters,Panulirus argus

Assessing the nutritional condition of field‐caught animals has obvious ecological utility, but few indices of condition have been developed for spiny lobsters. We evaluated two potential indices of nutritional condition for the Caribbean spiny lobster, Panulirus argus, using early benthic stage lobsters maintained in the laboratory on two food treatments (100% and 25% of maximum daily consumption) for four months. Every two weeks, we sampled lobsters to ascertain their condition using both a weight/carapace length ratio and triacylglycerol/bodyweight ratio. Triacylglycerol (TAG) content in the hepatopancreas, abdominal, and periopod tissue was determined using thin‐layer chromatography‐flame ionization detection. There was a significant difference in both carapace length and weight of lobsters in the two experimental treatments through time, confirming that our feeding treatments created a detectable difference in growth. We also found a corresponding significant difference between treatments for the weight/carapace length ratio, but the TAG/bodyweight ratio did not differ between treatments, and the variance among individuals was high.     

The olfactory pathway mediates sheltering behavior of Caribbean spiny lobsters, <Emphasis Type="Italic">Panulirus </Emphasis><Emphasis Type="Italic">argus</Emphasis>, to conspecific urine signals

The “noses” of diverse taxa are organized into different subsystems whose functions are often not well understood. The “nose” of decapod crustaceans is organized into two parallel pathways that originate in different populations of antennular sensilla and project to specific neuropils in the brain—the aesthetasc/olfactory lobe pathway and the non-aesthetasc/lateral antennular neuropil pathway. In this study, we investigated the role of these pathways in mediating shelter selection of Caribbean spiny lobsters, Panulirus argus, in response to conspecific urine signals. We compared the behavior of ablated animals and intact controls. Our results show that control and non-aesthetasc ablated lobsters have a significant overall preference for shelters emanating urine over control shelters. Thus the non-aesthetasc pathway does not play a critical role in shelter selection. In contrast, spiny lobsters with aesthetascs ablated did not show a preference for either shelter, suggesting that the aesthetasc/olfactory pathway is important for processing social odors. Our results show a difference in the function of these dual chemosensory pathways in responding to social cues, with the aesthetasc/olfactory lobe pathway playing a major role. We discuss our results in the context of why the noses of many animals contain multiple parallel chemosensory systems.     

In Memoriam: Professor Walter James Harman, PhD (1928–2002)

The Mediterranean spiny lobsters, the common spiny lobster Palinurus elephas (Fabricius, 1758) and the pink spiny lobster P. mauritanicus Gruvel, 1911, are important target species for commercial fisheries. In this study, we focus our attention on the DNA sequence variation of the mitochondrial cytochrome oxydase I gene (COI) in the two species of Palinurus. Spiny lobster DNA samples from four Mediterranean localities were analysed to examine the genetic variability at both the intra- and interspecific level. Furthermore, the phylogenetic relationships within the family Palinuridae (among the two species of Palinurus, most of the species of Panulirus and all the species of Jasus) are examined.     

An electromagnetic tracking system for studying the movements of rock (spiny) lobsters

A simple and inexpensive method for examining the movements of rock (spiny) lobsters is described. Electromagnetic tags emitting a pulsed signal were detected in loop antennae placed on the bottom in a grid pattern within a shallow coastal reef area. Signals were received in a tuned receiver and then interpreted using an oscilloscope. Results showing examples of the nocturnal movements of Panuliruscygnus George the western rock lobster in Western Australia are presented.     

Ontogenetic shifts in resource allocation: colour change and allometric growth of defensive and reproductive structures in the Caribbean spiny lobster Panulirus argus

Resource allocation theory predicts a disproportionately large allocation of resources to defensive structures during early ontogeny in organisms that are subject to more intense predation at smaller than at larger body sizes. We tested this prediction on the Caribbean spiny lobster Panulirus argus, which exhibits a negative relationship between predation risk and body size with a high natural mortality of smaller individuals. Independent allometric growth analyses demonstrated that numerous defensive structures (e.g. orbital horns, segments supporting the antenna, the tail fan) display negative allometric growth throughout ontogeny. We interpret these findings as lobsters investing disproportionately more resources to defensive structures when small to improve survivorship. Similarly, we observed an ontogenetic shift in lobster colour pattern; small individuals (< 23 mm carapace length) that inhabit nursery grounds (preferably among red algae) displayed a disruptive pattern (camouflage), whereas larger juveniles displayed a bicolour pigmentation typical of adult lobsters. This shift in colour pattern further suggests that small lobsters employ cryptic coloration throughout their asocial algal stage. However, this cryptic coloration offers no advantage when lobsters grow larger and start dwelling in crevices. Other structures linked to reproduction (e.g. female pleopods and male pereopods) experienced either isometric or positive allometric growth throughout ontogeny. Our results support one of the main predictions of resource allocation theory and demonstrate ontogenetic shifts in defensive structures and coloration concomitantly with changes in lobster mortality risk mediated by size‐dependent predation risk. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ●● , ●●–●●.     

Function of an eyestalk ganglion,the Medulla terminalis,in olfactory integration in the lobster,Panulirus argus

Summary Ipsilateral antennular dysfunction resulting from total unilateral eyestalk ablation in spiny lobsters does not occur when visual input is restricted by an opaque cap over one eyestalk, or when optic ganglia alone (eg. lamina ganglionaris, medulla externa, medulla interna) are removed. Antennular dysfunction appears only when connections between the most proximal of the four eyestalk ganglia, the medulla terminalis, and the remainder of the cerebral ganglia (brain) are interrupted. We conclude that neural processing of olfactory input from the antennule involves structures in the medulla terminalis.Contribution number 430 from the Bermuda Biological Station for Research, Inc. This work was supported by USPHS Grant NB-06017.     

The Constructional Properties of the Exoskeleton of Homarid, Palinurid, and Scyllarid Lobsters

Lobsters, as members of the Arthropoda, are already endowed with a laminated exoskeleton due to the mineralization of their cuticle. Mineralized laminate structures are found throughout animal phyla and convey, through their multiple surfaces, matrix planes that act as crack-blunting mechanisms. In addition, spiny and slipper lobsters, but not clawed lobsters, add a surface tubercle system that reflects a ventral surface pit system in the carapace. The architecture of these systems coincides with known strategies for crack blunting in composite materials. The division of vertical and horizontal crack blunting systems corresponding to laminate and tubercular systems and pit systems, respectively, may not be so easily separable in a functional sense. Although there is overlap in crack-blunting ability in both systems, separation into horizontal and vertical crack-blunting systems is a convenient way to discuss how skeletons of lobsters resist failure. While laminate structures dissipate forces horizontally in each layer, tubercle and pit systems serve to increase the surface area available to dissipate forces. These systems may represent evolutionary solutions to predation, particularly by predators that strike, crush or bite holes, rather than those that engulf.     

Species identification of spiny lobster phyllosome larvae via ribosomal DNA analysis.

Within the tropical northwestern Atlantic, Panulirus argus, P. guttatus, and P. laevicauda (Palinuridae family), are sympatric. Numerous studies have examined the distribution and abundance of planktonic phyllosome larvae with respect to recruitment of spiny lobsters to the benthic population, but the data are of limited use because larvae of these species cannot yet be distinguished from one another by morphological characteristics. A simple molecular method that unambiguously differentiates adults or larvae of P. argus, P. guttatus, and P. laevicauda is described: a 5' region of 28s ribosomal DNA is amplified in vitro and then cut with a diagnostic restriction enzyme to identify each species. Data are also presented from the application of this method to representative plankton tows.     

Diseases of spiny lobsters: a review

Spiny lobsters have few reported pathogens, parasites and symbionts. However, they do have a diverse fauna comprised of a pathogenic virus, several bacteria, protozoans, helminths and even symbiotic crustaceans. A few idiopathic syndromes have also been reported, but these appear correlated with lobsters held in poor conditions. Fungal and bacterial pathogens present significant threats for rearing spiny lobsters in aquaculture settings, but only one pathogen, Panulirus argus virus 1, is thought to have damaged a fishery for a spiny lobster. No doubt others will emerge as lobsters are brought into aquaculture setting and as fishing pressure intensifies with stocks become more susceptible to anthropogenic stressors.     

Identification of steroid hormones and fatty acids during gonadal maturation of spiny lobster Panulirus homarus

Information on steroid hormones and fatty acids that play roles in lobster reproduction is still very limited although the data are indispensable to seed production in hatchery. The study was designed to identify steroid hormones and fatty acids during gonadal maturation of spiny lobster (Panulirus homarus). Male spiny lobsters were treated with and without thyroxine injection. Female spiny lobsters were treated with and without eyestalk ablations during mature and immature gonad developments. Androst-5-en-17-one,3β (androst) and estran-3-one,17β (estran), two steroid hormones were identified at different levels of gonadal maturity of spiny lobsters. High concentrations of androst and estran were detected in the male spiny lobsters treated with thyroxine injections. Estran showed high concentrations in female brood stock of spiny lobsters during oogenesis stages both without eyestalk ablation and with ablation of one or two eyestalks, except in the immature female gonads. It was found that stearic acid was the highest and dominant fatty acid in mature male spiny lobster. Stearic acid, oleic acid, palmitic acid and caprylic acid were fatty acids with high concentrations in immature and mature female spiny lobsters. After 30 days in captivity, only stearic acid and oleic acid were found dominantly in eyestalk ablated mature female spiny lobsters.     

Subtidal-intertidal trophic links: American lobsters [Homarus americanus (Milne-Edwards)] forage in the intertidal zone on nocturnal high tides

Homarus americanus (Milne-Edwards), the American lobster, is a predator in New England subtidal communities, feeding on ecologically important grazers (sea urchins), mesopredators (crabs), and basal species (mussels). In this study, we provide the first report of adult American lobsters foraging in rocky intertidal habitats during nocturnal high tides. Censuses by SCUBA divers in the low intertidal (Chondrus crispus Stackhouse) zone showed mean densities of 2.2 lobsters/20 m² on nocturnal high tides, with contrasting low densities of 0.18/20 m² during diurnal high tides. Nocturnal high-tide intertidal densities were 62% of those reported in a previous study of lobsters in nearby subtidal rocky areas (Novak, 2004). The average carapace length of lobsters in the intertidal at night was > 50 mm. These lobsters were actively foraging in the intertidal with collected individuals having a mean stomach fullness of 67%. Prey found in the stomach contents primarily consisted of crabs, mussels and snails. Field experiments showed that lobsters rarely fed on medium to large size individuals of the common intertidal snail, Littorina littorea (L.). In contrast, experiments with local crab species demonstrated that lobsters actively and readily prey on Cancer irroratus (Say) and Carcinus maenas (L.), but were significantly less likely to consume Cancer borealis (Stimpson). The abundance of Carcinus maenas and blue mussels (Mytilus edulis L.) in the intertidal zone may explain the upshore movement of lobsters. Since nocturnal migration of Homarus americanus into the intertidal zone has not been documented before, our understanding of the dynamics of New England intertidal communities needs to be expanded to include this predator.     

Are Trade-Offs Among Species’ Ecological Interactions Scale Dependent? A Test Using Pitcher-Plant Inquiline Species

Trade-offs among species' ecological interactions is a pervasive explanation for species coexistence. The traits associated with trade-offs are typically measured to mechanistically explain species coexistence at a single spatial scale. However, species potentially interact at multiple scales and this may be reflected in the traits among coexisting species. I quantified species' ecological traits associated with the trade-offs expected at both local (competitive ability and predator tolerance) and regional (competitive ability and colonization rate) community scales. The most common species (four protozoa and a rotifer) from the middle trophic level of a pitcher plant (Sarracenia purpurea) inquiline community were used to link species traits to previously observed patterns of species diversity and abundance. Traits associated with trade-offs (competitive ability, predator tolerance, and colonization rate) and other ecological traits (size, growth rate, and carrying capacity) were measured for each of the focal species. Traits were correlated with one another with a negative relationship indicative of a trade-off. Protozoan and rotifer species exhibited a negative relationship between competitive ability and predator tolerance, indicative of coexistence at the local community scale. There was no relationship between competitive ability and colonization rate. Size, growth rate, and carrying capacity were correlated with each other and the trade-off traits: Size was related to both competitive ability and predator tolerance, but growth rate and carrying capacity were correlated with predator tolerance. When partial correlations were conducted controlling for size, growth rate and carrying capacity, the trade-offs largely disappeared. These results imply that body size is the trait that provides the basis for ecological interactions and trade-offs. Altogether, this study showed that the examination of species' traits in the context of coexistence at different scales can contribute to our understanding of the mechanisms underlying community structure.     

Agonistic interactions between the invasive green crab, Carcinus maenas (Linnaeus) and juvenile American lobster, Homarus americanus (Milne Edwards)

Invasive organisms have the potential for competition with native organisms. In the Southern Gulf of St. Lawrence, juvenile American lobsters have a potential spatial overlap with adult green crabs. Crustaceans use agonistic behaviour to settle disputes, with the larger organism often winning contests for limited resources such as food and shelter. Two experiments were carried out using adult green crabs (53-76 mm carapace width) and juvenile American lobsters (28-57 mm carapace length). The first experiment used a limited food resource. We found that green crabs were the first to the food in all trials, fed in significantly more trials than lobsters and spent a significantly greater proportion of time with the food. The lobsters were only able to displace the green crabs from the food in 2 of 65 attempts. The second experiment was designed to examine shelter competition; unexpectedly some predation by green crabs on lobsters occurred, which allowed us to test hypotheses about how relative size and shelter use affect predation. Green crabs captured and consumed juvenile lobsters in 6 of 11 trials. The lobsters that survived spent significantly more time in shelter. There was no clear relationship between shelter use and size of lobster. The lobsters that were larger in relation to the green crabs suffered a higher rate of predation, which we believe was due to more conspicuous activity and less use of shelter. It appears that green crabs have the potential to negatively impact native juvenile lobster.     

Calling Behavior and Morphology of the Stridulatory Apparatus in Neotropical Grasshoppers (Aleuas Stål; Orthoptera)

The sound production and sound producing apparatus of two species of grasshoppers, A. lineatus Stål, 1878 and A. vitticollis Stål, 1878 (Orthoptera, Acrididae, Copiocerinae) have been studied. For both species oscillograms and other physical data on sounds are presented for the first time. The sound producing behavior is described and the stridulation apparatus mechanism is illustrated. In the case of A. vitticollis these data are the first known on its sound production and general acoustic behavior.     

Zur Deutung der Carapaxfurchen der Astacidea (Crustacea, Decapoda)

The carapace grooves of decapod Crustacea are hitherto considered as remnants of somit boundaries. This hypothesis is rejected for theoretical reasons and on the basis of morphological studies of recent crayfishes and lobsters. The grooves are recognized as mechanically induced secondary structures resulting from the attachment of different muscles to the carapace. The grooves can be used for the diagnosis of certain systematic groups, but are of little value for phylogenetic considerations.     

The stomatopod rumble: Low frequency sound production in Hemisquilla californiensis

Stomatopods (mantis shrimp), numbering over 450 species, are renowned for their exceptional visual and chemosensory abilities and yet virtually nothing is known about their use of acoustic signals. We present acoustic recordings and analyses of the sounds of a stomatopod, Hemisquilla californiensis. This species generates tonal, low frequency sounds, which we term 'rumble', that are spectrally similar to those produced by African and Asian elephants. The fundamental frequency of the stomatopod rumble ranges from 20 to 60 Hz, with a strong second harmonic. Hemisquilla californiensis appears to generate these sounds through contractions of the posterior mandibular remotor muscle which is coupled to a stiff, lateral extension of the carapace. The sides of the carapace are covered by large, polarized, red spots which vibrate during sound production. Thus, the animals may generate a multi-modal signal by coupling vibrational and visual signals. Hemisquilla californiensis generates the rumble while interacting with potential predators and burrow intruders, suggesting a defensive or territorial function for the sound.     

Sea hares use novel antipredatory chemical defenses

Numerous studies have demonstrated that chemical defenses protect prey from predation and have often assumed that these defenses function by repelling predators. Surprisingly, few have investigated the mechanisms whereby predators are affected by these defenses. Here, we examine mechanisms of chemical defense of sea hares (Aplysia californica), which, when attacked by spiny lobsters (Panulirus interruptus), release defensive secretions from ink and opaline glands. We show that ink-opaline facilitates the escape of sea hares by acting through a combination of novel and conventional mechanisms. Ink-opaline contains millimolar quantities of amino acids that stimulate chemoreceptor neurons in the spiny lobster's nervous system. Ink stimulates appetitive and ingestive behavior, opaline can elicit appetitive behavior but can also inhibit ingestion and evoke escape responses, and both stimulate grooming. These results suggest that these secretions function by "phagomimicry," in which ink-opaline stimulates the feeding pathway to deceive spiny lobsters into attending to a false food stimulus, and by sensory disruption, in which the sticky and potent secretions cause high-amplitude, long-lasting chemo-mechanosensory stimulation. In addition, opaline contains a chemical deterrent that opposes appetitive effects. Thus, chemical defenses may act in more complex manners than palatability assays of prey chemistry may suggest.