Morphology of grooming limbs in species of Petrolisthes and Pachycheles (Crustacea: Decapoda: Anomura: Porcellanidae): a scanning electron microscopy study

The morphology of the grooming limbs, the 5th pair of pereiopods, was studied by scanning electron microscopy in six species of porcellanid crabs, Petrolisthes cabrilloi, P. cinctipes, P. armatus, P. galathinus, Pachycheles monilifer, and Pachycheles rudis, and their function was inferred by comparison with findings from previous studies. Grooming limb morphology was almost identical among the four Petrolisthes species and differed little compared to that of the two Pachycheles species. The 5th pereiopods bore a basal tuft of mechanoreceptive setae, three different types of grooming setae armed with setules or denticles, two types of smooth sensilla for location and identification of fouling objects, and a terminal, toothed chela for picking firmly attached objects off the gills and body. The grooming limb was extremely flexible and could reach most parts of the body, including the gill chamber on the opposite side. The grooming limb morphology in Petrolisthes cabrilloi is consistent with its wellknown effectiveness in preventing parasitism by the rhizocephalan Lernaeodiscus porcellanae. Grooming setae remove recently attached cyprids, while the chela can grip and remove the much smaller, firmly attached kentrogons. Porcellanid crabs not known to host rhizocephalans, however, had grooming limbs almost identical to those of Petrolisthes cabrilloi despite their previously demonstrated failure to prevent settlement and infestation by L. porcellanae larvae. The effectiveness of P. cabrilloi in removing kentrogons, therefore, seems also to depend on behavioral adaptations whereby this species recognizes the parasite larvae as high-threat objects.     

Pheromones and colonization: reassessment of the milkweed bug migration model (Heteroptera: Lygaeidae: Lygaeinae)

Summary. Research on insect migration has justifiably emphasized females – the so-called “oogenesis-flight syndrome”– since it is the females that place the eggs into new habitats. The large and small milkweed bugs, Oncopeltus fasciatus and Lygaeus kalmii, respectively, have featured prominently in studies of insect migration and sequestration of host plant toxins for chemical defense. Here we report that males of these species, and males of another well-studied lygaeine (Neacoryphus bicrucis), produce pheromones in glands usually considered to serve only a defensive role in Heteroptera (the metathoracic scent glands), and that these pheromones are exploited by a tachinid parasitoid as a host-finding kairomone. The pheromones are mixtures of C₆ and C₈ saturated and unsaturated esters reminiscent of lepidopteran pheromones, and the key compound of the O. fasciatus pheromone has now been correctly identified as (E)-2,7-octadienyl acetate. It is proposed that the concept of the oogenesis-flight syndrome for these kinds of insects should accommodate the role of males in the migration process. The hypothesis is presented that male-produced pheromones play a significant role in guiding colonization of new habitats in many heteropteran species. In addition, data are presented suggesting that there is a trade-off between the amount of pheromone produced by colonizing males and the host breadth of the species. Received 21 December 1998; accepted 15 February 1999.     

The behaviour of giant clams (Bivalvia: Cardiidae: Tridacninae)

Giant clams, the largest living bivalves, live in close association with coral reefs throughout the Indo-Pacific. These iconic invertebrates perform numerous important ecological roles as well as serve as flagship species—drawing attention to the ongoing destruction of coral reefs and their associated biodiversity. To date, no review of giant clams has focussed on their behaviour, yet this component of their autecology is critical to their life history and hence conservation. Almost 100 articles published between 1865 and 2014 include behavioural observations, and these have been collated and synthesised into five sections: spawning, locomotion, feeding, anti-predation, and stress responses. Even though the exact cues for spawning in the wild have yet to be elucidated, giant clams appear to display diel and lunar periodicities in reproduction, and for some species, peak breeding seasons have been established. Perhaps surprisingly, giant clams have considerable mobility, ranging from swimming and gliding as larvae to crawling in juveniles and adults. Chemotaxis and geotaxis have been established, but giant clams are not phototactic. At least one species exhibits clumping behaviour, which may enhance physical stabilisation, facilitate reproduction, or provide protection from predators. Giant clams undergo several shifts in their mode of acquiring nutrition; starting with a lecithotrophic and planktotrophic diet as larvae, switching to pedal feeding after metamorphosis followed by the transition to a dual mode of filter feeding and phototrophy once symbiosis with zooxanthellae (Symbiodinium spp.) is established. Because of their shell weight and/or byssal attachment, adult giant clams are unable to escape rapidly from threats using locomotion. Instead, they exhibit a suite of visually mediated anti-predation behaviours that include sudden contraction of the mantle, valve adduction, and squirting of water. Knowledge on the behaviour of giant clams will benefit conservation and restocking efforts and help fine-tune mariculture techniques. Understanding the repertoire of giant clam behaviours will also facilitate the prediction of threshold levels for sustainable exploitation as well as recovery rates of depleted clam populations.     

Patterns of trace-metal distribution in tissues of Pleoticus muelleri (Crustacea: Decapoda: Solenoceridae)

Trace-metal distribution in tissues of the shrimp Pleoticus muelleri Bate from the Patagonian region, Argentina, was related to sex, size and physiological condition. Concentrations of cadmium, copper, manganese and zinc were determined in the digestive gland, male reproductive system and muscle of adult specimens. Significant (p<0.01) sex-dependent differences in mean metal concentrations (g g^-1 wet wt) were found in the following tissues of males and females, respectively: digestive gland, Cu, 82.9 and 30.8; Zn, 32.5 and 44.9; reproductive system, Zn, 12.8 and 38.6; Mn, 1.7 and 3.1; Cd, 0.29 and 0.58. Metal levels of muscle showed no significant differences between sexes. Cadmium was not detected in muscle, suggesting that its incorporation into this tissue was strictly regulated. Metal concentrations in the male reproductive system were in general weakly correlated (0.001<p<0.05) with either body size or the concentration of metals in the digestive gland. The developing ovary incorporated substantial amounts of zinc, while cadmium and copper levels decreased at proportional rates (p<0.001). The patterns of the variations in the concentrations of these metals in the ovary strongly suggested that a regulatory mechanism operated throughout oogenesis. Manganese was not involved in this presumptive mechanism. The results of this study are discussed in terms of concentrations and in absolute amounts for standardized individuals.     

Podotreme affinities of Raninoides sp. and Lyreidus brevifrons: evidence from spermatozoal ultrastructure (Crustacea: Brachyura: Raninoidea)

Spermatozoal synapomorphies which singly or collectively distinguish Raninoidea are: (1) the presence of single (Ranina, Raninoides) or multiple (Lyreidus) keel-like projections of the acrosomal capsule; (2) a very large, weakly electron-opaque peripheral acrosomal zone (Ranina, Raninoides) and an homologous large outer zone in Lyreidus; (3) poor differentiation of the operculum from the capsule (autapomorphy); (4) a very well developed, perforate subopercular zone, of variable form; (5) presence of unique inward longitudinal projections (septa or corrugations) in the wall of the subacrosomal chamber (autapomorphy). Shared, presumably synapomorphic characters of Ranina and Raninoides but not of Lyreidus within the Raninidae, are: (1) branching of some of the subacrosomal septa (unbranched in Lyreidus); (2) the subspheroidal form of the acrosome in Ranina with a length: width ratio (L:W) of 0.76, and, although slightly more depressed, in Raninoides (L:W 0.73), considered apomorphic relative to the more depressed form in Lyreidus (L:W 0.52); (3) single or multiple coiled perforatorial filaments (Ranina, Raninoides) contrasting with a capitate perforatorium with “amoeboid” head in Lyreidus; (4) division of the capsule wall to give one posterior (Ranina) or multiple enclaves, plesiomorphically (?) absent in Lyreidus. Similarities of Lyreidus with other podotremes include the capitate perforatorium, questionably related to the radiate spiked-wheel structure of homolids in which acrosome proportions are similar or less closely to the bilateral capitate perforatorium of dromiids and dynomenids, and basal capsular projections as in the dromiid Stimdromia (=Petamolera) lateralis and in cyclodorippoids. No spermatozoal synapomorphies support a sister-group relationship of raninoids and heterotreme-thoracotreme crabs.     

Anatomy of structures associated with air-breathing in Orchestia gammarellus (Crustacea: Amphipoda: Talitridae): coxal plates and gills

The structure of the coxal gills and coxal plates of the semi-terrestrial beachflea Orchestia gammarellus (Pallas) (Crustacea: Amphipoda: Talitridae) is described in relation to their possible use for aerial gas exchange and ion exchange. Anatomical evidence is presented to support the hypothesis that the medial surface of the coxal plates functions as an extrabranchial aerial gas-exchange site in O. gammarellus. Thus, the effective diffusion distance across the medial (or inside-facing) surface of O. gammarellus coxal plates (mean±SD=5.4±0.3 m; n=9, cuticle thickness 4.4±0.5 m, n=21) is only a third of the equivalent distance across both the coxal gills (18.4±6.0 m, n=10; cuticle thickness 1.7±0.6 m, n=7) and the lateral (or external) surface of the coxal plates (19.4±0.7 m, n=5; cuticle thickness 8.7±0.8 m, n=7). Chloride-ion-permeable areas were located using a silver-staining technique. All ten coxal gills appeared to be equally permeable to chloride ions after examination with a light microscope. However, the coxal plates and the rest of the integument do not appear to be chloride-permeable.     

Correction to: Contrasting effects of engineered carbon nanotubes on plants: a review

Environmental Geochemistry and Health - Unfortunately, in the original publication of the article, Prof. Yong Sik Ok’s affiliation was incorrectly published. The author’s affiliation is...     

Post-settlement mortality of juvenile lagoonal cockles (Cerastoderma glaucum : Mollusca: Bivalvia)

Like many benthic marine invertebrates the lagoonal cockle Cerastoderma glaucum Bruguière (Mollusca: Bivalvia) suffers extensive mortality between settling out of the plankton and reaching adolescence. It is thought that predation could be a major cause of this mortality. Experiments were conducted in aquaria to assess the influence of predation on the survival of juvenile lagoonal cockles. Newly settled spat (collected from Holkham Salts Hole, a lagoon in North Norfolk) were enclosed with potential predators at field densities. Predators used were fish, prawns and polychaetes (singly or combination). The numbers and sizes of the surviving cockles were measured after a 4-week period to establish the extent of predation, and the results were consistent with predation by the fish (Pomatoschistus microps) being a major causative agent of the mortality seen in the field. This reinforces the idea of epibenthic predation as an important structuring element for marine benthic communities. The effect of post-settlement juvenile mortality on cockle population demographics is considered, and other possible causes of mortality in the field are discussed. Received: 1 February 1998 / Accepted: 2 April 1998     

Feeding responses of the tentaculate deposit-feederEupolymnia nebulosa (Annelida: Polychaeta): Influence of sexual maturity

Feeding responses of the tentaculate depositfeeding polychaeteEupolymnia nebulosa (Montagu) were studied by measuring rates of uptake of three different¹⁴C-labelled diatoms (unialgal cultures ofNavicula incerta Grunow,Nitzschia acicularis Wm Smith, andNitzschia sp.). Worms used during this study were collected in the harbor of Port-Vendre (Western Mediterranean) during August 1986 (immature worms) and December 1987 (mature worms). Uptake rates were affected both by the length of the experiments and by the nature of the food offered. The highest rate of uptake (2.98 10^–4 mg ashfree dry wt of algae mg^–1 dry wt of worms h^–1) was obtained during short-term experiments (4 h) with the smallest diatom (Nitzschia sp.). The lowest rate of uptake (0.21 10^–4 mg ash-free dry wt of algae mg^–1 dry wt of worms h^–1) was also obtained withNitzschia sp., but for a long-term (48 h) experiment. There was no significant difference between rates of uptake of immature and mature worms.     

Antifouling adaptations of marine shrimp (Crustacea: Decapoda: Caridea): Functional morphology and adaptive significance of antennular preening by the third maxillipeds

Preening (cleaning, grooming) of the antennules and other cephalothoracic appendages by the third maxillipeds was observed in several species of shrimp. Distribution and ultrastructure of serrate grooming setae on the third maxillipeds, which scrape the antennules, was studied with light and scanning electron microscopy. The motor patterns of antennular cleaning were similar for all species. Antennular preening was the most frequent grooming behavior observed, but the duration of other grooming behaviors was greater. Tidepool shrimps (Heptacarpus pictus) experimentally prevented from grooming the antennules by ablation suffered fouling of the olfactory hairs of the antennules with their subsequent breakage and loss; antennules of controls remained clean and undamaged. Antennular preening, a frequent and widespread behavior of caridean shrimps and other decapod crustaceans, is suggested as having high adaptive value in keeping sensory sites free of epizoic and sedimentary fouling which might render them inoperative.     

Ultrastructure of the mature spermatozoon of the coconut crabBirgus latro (Coenobitidae: Paguroidea: Decapoda)

The spermatozoon ofBirgus latro (Linnaeus, 1767) is approx 14µm in length. It is composed of a large multi-layered oblong-ovoid acrosome which is capped by a conical operculum and lies anterior to a small ring of cytoplasm and an amorphous nucleus which is drawn out into a series of arms or extensions. Originating from the cytoplasmic area are three further long microtubular arms. The sperm ofB. latro is very similar to the sperm of the only other genus in the Coenobitidae,Coenobita, of whichC. clypeatus is a representative species. They share a suite of ultrastructural characters including: a long, cylindrical, capsule-bound acrosome containing an inner acrosome core, a large acrosome ray zone and a thin outer acrosome zone; an apical operculum anterior to a subopercular zone divided into two areas of differing density; an invaginated perforatorial zone with a bipartite granular matrix; microvillus-like extensions of the inner acrosome core projecting into the perforatorial invagination; a ring of cytoplasm, around the base of the acrosome, containing numerous mitochondria, extensive lamellar systems and the bases of three microtubular arms; granular nuclear material forming irregular arms; and at the posterior portion of the cell membrane a combination of nuclear and plasma membranes. Some ultrastructural characters which separate the two genera are: a domed operculum inC. clypeatus as opposed to a conical one inB. latro; inB. latro there is some residual cytoplasm external to the operculum and centrioles are absent from the mature spermatozoon; inC. clypeatus the inner acrosome core does not appear to invest the perforatorium and a series of dense rods are found subjacent and internal to the operculum. Similarities between the two coenobitids are greater than those shared with the paguridEupagurus bernhardus. In this latter species, the acrosome is more ovoid than cylindrical and the acrosome zones are less conspicuous; the operculum is absent from the mature sperm (although present during spermiohistogenesis); no subopercular zone is present; the perforatorium contains longitudinally arranged microtubules and extends the full length of the acrosome; and the nuclear material does not form separate nuclear arms. Sperm ultrastructure supports monophyly of the Paguroidea, while distinguishing coenobitids from pagurids within this superfamily.     

Predator detection and avoidance: Ornamentation of tube-caps of Diopatra spp. (Polychaeta: Onuphidae)

One strategy for predator avoidance involves the creation of a structural refuge. Onuphid polychaetes characteristically ornament above-sediment portions of their tubes (=tube-caps) with shell and algal debris. These species feed on the sediment surface through an opening in the tube-cap, and thus the ability to detect a surface predator while feeding would be advantageous. An investigation of the function of the ornamentation in Diopatra spp. suggests that ornamentation facilitates predator detection and avoidance. Three intensities of mechanical disturbance were applied directly to D. ornata tube-caps. When ornamented tube-caps were stimulated, the response of worms to the three intensities were significantly different, and increased in duration with greater intensities. In contrast, when no ornamentation was present, the responses were not significantly different, and were similar to the low-intensity response when ornamentation was present. This suggests that ornamentation should allow a worm to distinguish between harmful (high intensity due to mobile epifaunal predators) and profitable (low intensity due to drift algae) disturbances, and furthermore, worms with ornamented tube-caps should be more successful in escaping surface predators. Densities of intertidal populations of D. cuprea at Tom's Cove, Virginia, USA, correlated with the amount of tube ornamentation, consistent with this predator detection and avoidance hypothesis. Final tube-cap lengths of laboratory D. ornata and field D. cuprea were inversely related to the size of attached debris. When large debris was attached, cap formation ceased earlier and caps were shorter than when small debris or no debris was attached. Cryptic and food-catching functions would predict that highly ornamented tubes would be most advantageous, while only a few large debris would be required for disturbance transmission. Laboratory specimens showed no selectivity between 0.5 or 1.5 cm² shell; or 1.0 cm² or 3 to 8 cm² algae, and utilized shell and algae when available and according to relative abundance. Tube-caps of field specimens also showed positive correlations between shell attached and shell abundance in the local sediment. While such lack of selectivity may enhance cryptic properties of the tube-cap, it is argued that conditions seldom exist which would permit selectivity of debris size or of specific material. These data are consistent with the hypothesis that ornamentation functions as a created refuge for predator detection and avoidance and further suggest that the availability of ornamental debris in the environment indirectly can affect these species' distributions and abundances.     

Moderate stoichiometric homeostasis in the sea urchin Lytechinus variegatus: effects of diet and growth on C:N:P ratios

The influence of dietary elemental contents on consumer stoichiometry was investigated in selected and combined soft tissues (as a proxy of the whole individual) of the omnivorous sea urchin, Lytechinus variegatus. We raised urchins for 4 months in controlled seawater tanks using three different diets with different nutritional contents (from lower to higher: seagrass, red macroalgae, and a formulated diet). Individuals fed the different diets varied an average of 19.7, 19.4, and 38 % in C:N, C:P, and N:P ratios, respectively, with stronger temporal variability for C:P and N:P ratios across tissues and whole individuals. This resulted in homeostasis parameters (1/H) of ?0.45, 0.09, and 0.38, respectively, for C:N, C:P, and N:P, indicative of homeostatic to weakly homeostatic organisms, at least for C:P and N:P ratios. Individuals fed the nutrient-rich formulated diet had higher growth rates (14 ± 0.83 g WW month^?1) than those fed macroalgae or seagrass (9.3 ± 0.57 and 3.4 ± 0.33 g WW month^?1, respectively). However, rapid body increments in more nutritional diets caused both a decrease in the %N and an increase in the %P of soft tissues, which resulted in significant but opposite effects of diet stoichiometry and growth in sea urchin C:N (R = ?0.74 and R = 0.93, for diet and growth effects, respectively) and N:P ratios (R = 0.60 and R = ?0.63, also, respectively, for diet and growth effects). Among potential compensatory mechanisms helping to preserve certain levels of homeostasis, ingestion rates (g WW diet per g WW of urchin) were higher for seagrass and macroalgae diets than for the nutrient-rich formulated diet. In contrast, absorption and growth efficiencies displayed significant negative associations with nutrient contents in diets and did not exhibit nutritional compensation. Overall, our results suggest that resource stoichiometry strongly determines the growth rate of individuals (R = 0.88, P < 0.01), and moderate variability in C:N:P ratios of sea urchins possibly arise from differences in the allocation of proteins and RNA to body components, similarly to what has been proposed by the growth rate hypothesis.     

Correction to: Impact of biogas digestates on soil microbiota in agriculture: a review

Environmental Chemistry Letters -     

Spawning behaviour and egg capsules of Concholepas concholepas (Mollusca: Gastropoda: Muricidae)

Observations on the spawning behaviour of the commercially important muricid snail Concholepas concholepas are reported. The sequence of events relating to external transportation, molding and hardening, and attachment of the egg capsules to a substratum were timed and divided into 6 major events. The snails studied in the laboratory spawned between April and Septemer. These results and data from the literature reveal an all-year-round spawning activity for the species. The frequency, size and time of spawning were studied over 1 year. Spawning activity always began at night, suggesting that the species retains its spawning rhythm in the laboratory. These was a direct relationship between length of the snail and size of the capsules spawned. No nutritive eggs were found; the number per capsule varied between 668 and 14,250 according to capsule size. Hatching time varied between 69 and 128 days at 13.5° to 14.5°C.     

Reprint of: Linking conservation and welfare: A theoretical model with application to Nepal

Theoretical articles linking conservation and welfare find a negative relationship between these two variables while empirical studies show that land protection may be positively related to welfare. Several authors attribute this empirical result to the development of ecotourism in protected areas. We thus argue that the gap between the theory and existing empirical results is partly explained by the fact that most theoretical models do not account for a productive activity on protected land. Therefore, we develop a theoretical model in which conservation allows developing an alternative sector and show that the relationship between conservation and welfare is U-shaped. We test this theoretical prediction using Nepalese data and find that conservation combined with ecotourism is indeed positively related to local welfare.