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Concept: Crustacean


We investigated the navigational capabilities of the world’s largest land-living arthropod, the giant robber crab Birgus latro (Anomura, Coenobitidae); this crab reaches 4 kg in weight and can reach an age of up to 60 years. Populations are distributed over small Indo-Pacific islands of the tropics, including Christmas Island (Indian Ocean). Although this species has served as a crustacean model to explore anatomical, physiological, and ecological aspects of terrestrial adaptations, few behavioral analyses of it exist. We used a GPS-based telemetric system to analyze movements of freely roaming robber crabs, the first large-scale study of any arthropod using GPS technology to monitor behavior. Although female robber crabs are known to migrate to the coast for breeding, no such observations have been recorded for male animals. In total, we equipped 55 male robber crabs with GPS tags, successfully recording more than 1,500 crab days of activity, and followed some individual animals for as long as three months. Besides site fidelity with short-distance excursions, our data reveal long-distance movements (several kilometers) between the coast and the inland rainforest. These movements are likely related to mating, saltwater drinking and foraging. The tracking patterns indicate that crabs form route memories. Furthermore, translocation experiments show that robber crabs are capable of homing over large distances. We discuss if the search behavior induced in these experiments suggests path integration as another important navigation strategy.

Concepts: Crustacean, Coconut crab, Coenobitidae, Anomura, Christmas Island, Caroline Island, Chagos Archipelago


Carbonated hydroxyapatite is the mineral found in vertebrate bones and teeth, whereas invertebrates utilize calcium carbonate in their mineralized organs. In particular, stable amorphous calcium carbonate is found in many crustaceans. Here we report on an unusual, crystalline enamel-like apatite layer found in the mandibles of the arthropod Cherax quadricarinatus (freshwater crayfish). Despite their very different thermodynamic stabilities, amorphous calcium carbonate, amorphous calcium phosphate, calcite and fluorapatite coexist in well-defined functional layers in close proximity within the mandible. The softer amorphous minerals are found primarily in the bulk of the mandible whereas apatite, the harder and less soluble mineral, forms a wear-resistant, enamel-like coating of the molar tooth. Our findings suggest a unique case of convergent evolution, where similar functional challenges of mastication led to independent developments of structurally and mechanically similar, apatite-based layers in the teeth of genetically remote phyla: vertebrates and crustaceans.

Concepts: Arthropod, Calcium, Crustacean, Mineral, Calcium carbonate, Teeth, Crayfish, Aragonite


Biogenic amines, particularly serotonin, are recognised to play an important role in controlling the aggression of invertebrates, whereas the effect of neurohormones is still underexplored. The crustacean Hyperglycemic Hormone (cHH) is a multifunctional member of the eyestalk neuropeptide family. We expect that this neuropeptide influences aggression either directly, by controlling its expression, or indirectly, by mobilizing the energetic stores needed for the increased activity of an animal. Our study aims at testing such an influence and the possible reversion of hierarchies in the red swamp crayfish, Procambarus clarkii, as a model organism. Three types of pairs of similarly sized males were formed: (1) ‘control pairs’ (CP, n = 8): both individuals were injected with a phosphate saline solution (PBS); (2) ‘reinforced pairs’ (RP, n = 9): the alpha alone was injected with native cHH, and the beta with PBS; (3) ‘inverted pairs’ (IP, n = 9): the opposite of (2). We found that, independently of the crayfish’s prior social experience, cHH injections induced (i) the expression of dominance behaviour, (ii) higher glycemic levels, and (iii) lower time spent motionless. In CP and RP, fight intensity decreased with the establishment of dominance. On the contrary, in IP, betas became increasingly likely to initiate and escalate fights and, consequently, increased their dominance till a temporary reversal of the hierarchy. Our results demonstrate, for the first time, that, similarly to serotonin, cHH enhances individual aggression, up to reverse, although transitorily, the hierarchical rank. New research perspectives are thus opened in our intriguing effort of understanding the role of cHH in the modulation of agonistic behaviour in crustaceans.

Concepts: Arthropod, Hierarchy, Crustacean, Procambarus clarkii, Crayfish, Decapoda, Cambaridae, Procambarus


The amphipod crustacean Parhyale hawaiensis is a blossoming model system for studies of developmental mechanisms and more recently regeneration. We have sequenced the genome allowing annotation of all key signaling pathways, transcription factors, and non-coding RNAs that will enhance ongoing functional studies. Parhyale is a member of the Malacostraca clade, which includes crustacean food crop species. We analysed the immunity related genes of Parhyale as an important comparative system for these species, where immunity related aquaculture problems have increased as farming has intensified. We also find that Parhyale and other species within Multicrustacea contain the enzyme sets necessary to perform lignocellulose digestion (‘wood eating’), suggesting this ability may predate the diversification of this lineage. Our data provide an essential resource for further development of Parhyale as an experimental model. The first malacostracan genome will underpin ongoing comparative work in food crop species and research investigating lignocellulose as an energy source.

Concepts: DNA, Gene, Gene expression, RNA, Developmental biology, Model organism, Crustacean, Malacostraca


The American brine shrimp Artemia franciscana is invasive in the Mediterranean region where it has displaced native species (the sexual A. salina, and the clonal A. parthenogenetica) from many salt pond complexes. Artemia populations are parasitized by numerous avian cestodes whose effects have been studied in native species. We present a study from the Ebro Delta salterns (NE Spain), in a salt pond where both A. franciscana and native A. salina populations coexist, providing a unique opportunity to compare the parasite loads of the two sexual species in syntopy. The native species had consistently higher infection parameters, largely because the dominant cestode in A. salina adults and juveniles (Flamingolepis liguloides) was much rarer in A. franciscana. The most abundant cestodes in the alien species were Eurycestus avoceti (in adults) and Flamingolepis flamingo (in juveniles). The abundance of E. avoceti and F. liguloides was higher in the A. franciscana population syntopic with A. salina than in a population sampled at the same time in another pond where the native brine shrimp was absent, possibly because the native shrimp provides a better reservoir for parasite circulation. Infection by cestodes caused red colouration in adult and juvenile A. salina, and also led to castration in a high proportion of adult females. Both these effects were significantly stronger in the native host than in A. franciscana with the same parasite loads. However, for the first time, significant castration effects (for E. avoceti and F. liguloides) and colour change (for six cestode species) were observed in infected A. franciscana. Avian cestodes are likely to help A. franciscana outcompete native species. At the same time, they are likely to reduce the production of A. franciscana cysts in areas where they are harvested commercially.

Concepts: Arthropod, Crustacean, Brine shrimp, Transmission and infection of H5N1, Cestoda


Pig carcasses, as human proxies, were placed on the seabed at a depth of 300 m, in the Strait of Georgia and observed continuously by a remotely operated camera and instruments. Two carcasses were deployed in spring and two in fall utilizing Ocean Network Canada’s Victoria Experimental Network under the Sea (formerly VENUS) observatory. A trial experiment showed that bluntnose sixgill sharks could rapidly devour a carcass so a platform was designed which held two matched carcasses, one fully exposed, the other covered in a barred cage to protect it from sharks, while still allowing invertebrates and smaller vertebrates access. The carcasses were deployed under a frame which supported a video camera, and instruments which recorded oxygen, temperature, salinity, density, pressure, conductivity, sound speed and turbidity at per minute intervals. The spring exposed carcass was briefly fed upon by sharks, but they were inefficient feeders and lost interest after a few bites. Immediately after deployment, all carcasses, in both spring and fall, were very rapidly covered in vast numbers of lyssianassid amphipods. These skeletonized the carcasses by Day 3 in fall and Day 4 in spring. A dramatic, very localized drop in dissolved oxygen levels occurred in fall, exactly coinciding with the presence of the amphipods. Oxygen levels returned to normal once the amphipods dispersed. Either the physical presence of the amphipods or the sudden draw down of oxygen during their tenure, excluded other fauna. The amphipods fed from the inside out, removing the skin last. After the amphipods had receded, other fauna colonized such as spot shrimp and a few Dungeness crabs but by this time, all soft tissue had been removed. The amphipod activity caused major bioturbation in the local area and possible oxygen depletion. The spring deployment carcasses became covered in silt and a black film formed on them and on the silt above them whereas the fall bones remained uncovered and hence continued to be attractive to large numbers of spot shrimp. The carcass remains were recovered after 166 and 134 days respectively for further study.

Concepts: Oxygen, Arthropod, Water pollution, Crustacean, Malacostraca, Strait of Georgia, Dungeness crab, Bluntnose sixgill shark


It has been suggested that we do not know within an order of magnitude the number of all species on Earth [May RM (1988) Science 241(4872):1441-1449]. Roughly 1.5 million valid species of all organisms have been named and described [Costello MJ, Wilson S, Houlding B (2012) Syst Biol 61(5):871-883]. Given Kingdom Animalia numerically dominates this list and virtually all terrestrial vertebrates have been described, the question of how many terrestrial species exist is all but reduced to one of how many arthropod species there are. With beetles alone accounting for about 40% of all described arthropod species, the truly pertinent question is how many beetle species exist. Here we present four new and independent estimates of beetle species richness, which produce a mean estimate of 1.5 million beetle species. We argue that the surprisingly narrow range (0.9-2.1 million) of these four autonomous estimates-derived from host-specificity relationships, ratios with other taxa, plant:beetle ratios, and a completely novel body-size approach-represents a major advance in honing in on the richness of this most significant taxon, and is thus of considerable importance to the debate on how many species exist. Using analogous approaches, we also produce independent estimates for all insects, mean: 5.5 million species (range 2.6-7.8 million), and for terrestrial arthropods, mean: 6.8 million species (range 5.9-7.8 million), which suggest that estimates for the world’s insects and their relatives are narrowing considerably.

Concepts: Organism, Insect, Fungus, Arthropod, Animal, Crustacean, Beetle, Spider


The nervous system provides a fundamental source of data for understanding the evolutionary relationships between major arthropod groups. Fossil arthropods rarely preserve neural tissue. As a result, inferring sensory and motor attributes of Cambrian taxa has been limited to interpreting external features, such as compound eyes or sensilla decorating appendages, and early-diverging arthropods have scarcely been analysed in the context of nervous system evolution. Here we report exceptional preservation of the brain and optic lobes of a stem-group arthropod from 520 million years ago (Myr ago), Fuxianhuia protensa, exhibiting the most compelling neuroanatomy known from the Cambrian. The protocerebrum of Fuxianhuia is supplied by optic lobes evidencing traces of three nested optic centres serving forward-viewing eyes. Nerves from uniramous antennae define the deutocerebrum, and a stout pair of more caudal nerves indicates a contiguous tritocerebral component. Fuxianhuia shares a tripartite pre-stomodeal brain and nested optic neuropils with extant Malacostraca and Insecta, demonstrating that these characters were present in some of the earliest derived arthropods. The brain of Fuxianhuia impacts molecular analyses that advocate either a branchiopod-like ancestor of Hexapoda or remipedes and possibly cephalocarids as sister groups of Hexapoda. Resolving arguments about whether the simple brain of a branchiopod approximates an ancestral insect brain or whether it is the result of secondary simplification has until now been hindered by lack of fossil evidence. The complex brain of Fuxianhuia accords with cladistic analyses on the basis of neural characters, suggesting that Branchiopoda derive from a malacostracan-like ancestor but underwent evolutionary reduction and character reversal of brain centres that are common to hexapods and malacostracans. The early origin of sophisticated brains provides a probable driver for versatile visual behaviours, a view that accords with compound eyes from the early Cambrian that were, in size and resolution, equal to those of modern insects and malacostracans.

Concepts: Central nervous system, Nervous system, Brain, Insect, Arthropod, Human brain, Crustacean, Hexapoda


Pentastomids (tongue worms) are worm-like arthropods known today from ∼140 species [1]. All but four are parasitic on vertebrates. Their life cycle typically involves larval development in an intermediate host followed by maturation in the respiratory tract of a definitive terrestrial host. Fossil pentastomids are exceedingly rare and are known only from isolated juveniles [2-6]. The identity of the possible hosts of fossil pentastomids and the origin of their lifestyle have generated much debate. A new, exceptionally preserved species, described based on adults from 425-million-year-old marine rocks, is the only known fossil pentastomid associated with a host, in this case a species of ostracod crustacean. The pentastomids are preserved near eggs within the ostracod and also, uniquely for any fossil or living pentastomid, are attached externally to the host. This discovery affirms the origin of pentastomids as ectoparasitic on marine invertebrates. The terrestrialization of pentastomids may have occurred in parallel with the vertebrate invasion of land.

Concepts: Species, Insect, Crustacean, Vertebrate, Annelid, Invertebrate, Maxillopoda, Devonian


Small plastic detritus, termed ‘microplastics’, are a widespread and ubiquitous contaminant of marine ecosystems across the globe. Ingestion of microplastics by marine biota, including mussels, worms, fish and seabirds, has been widely reported, but despite their vital ecological role in marine food-webs, the impact of microplastics on zooplankton remains under-researched. Here, we show that microplastics are ingested by, and may impact upon, zooplankton. We used bio-imaging techniques to document ingestion, egestion and adherence of microplastics in a range of zooplankton common to the northeast Atlantic, and employed feeding rate studies to determine the impact of plastic detritus on algal ingestion rates in copepods. Using fluorescence and coherent anti-Stokes Raman scattering (CARS) microscopy we identified that thirteen zooplankton taxa had the capacity to ingest 1.7 - 30.6 µm polystyrene beads, with uptake varying by taxa, life-stage and bead-size. Post-ingestion, copepods egested faecal pellets laden with microplastics. We further observed microplastics adhered to the external carapace and appendages of exposed zooplankton. Exposure of the copepod Centropages typicus to natural assemblages of algae with and without microplastics showed that 7.3 µm microplastics (>4000 ml-1) significantly decreased algal feeding. Our findings imply that marine microplastic debris can negatively impact upon zooplankton function and health.

Concepts: Ecology, Eating, Ingestion, Coprophagia, Crustacean, Raman spectroscopy, Plankton, Copepod