Shell choice in<Emphasis Type="Italic"> Pagurus longicarpus</Emphasis> hermit crabs: does predation threat influence shell selection behavior?

Pagurus longicarpus hermit crabs depend on empty gastropod shells for protection against predation. Hermit crabs avoid gastropod shells in which holes have been drilled by naticid gastropods, and hermit crabs forced to occupy drilled shells are more vulnerable to predation by green crabs, Carcinus maenas. In this study, we examined the effect of predator cues on P. longicarpus shell investigation behavior and shell choice. In paired laboratory shell choice trials, we examined hermit crab response to green crab chemical cues. We compared hermit crabs from two sites differing in the percentage of Littorina littorea shells with drill holes. The percentage of time hermit crabs spent occupying intact shells increased significantly in the presence of predator cues. The effect of predator cues on the amount of time hermit crabs spent investigating shells differed between individuals from the two sites. Predator effluent had a marginal effect on the proportion of hermit crabs initially choosing intact shells and within 15 min most hermit crabs in both treatments occupied intact shells due to shell switching. These results indicate that predation cues alter P. longicarpus shell choice behavior favoring intact shells, which provide greater protection. In summary, predation appears to be a key factor influencing hermit crab shell selection behavior.Communicated by T. Czeschlik     

Competition for inorganic carbon between photosynthesis and calcification in the symbiont-bearing foraminifer Amphistegina lobifera

The feeding behaviour of the subtropical, intertidal crab Gaetice depressus (He Haan) has been investigated. This species was found to be an omnivorous scavenger capable of ingesting both animal and plant tissues which are passed to the mouth using the chelipeds. Furthermore, the mouth-parts themselves are modified and bear elongated setae, notably on the third maxillae. The net-like structures which the setae give rise to can be repeatedly flailed through the seawater in unison enabling the crabs to feed on suspended material. Beating frequency decreased with crab size but rose with increasing suspension concentration. Experiments demonstrated that suspension feeding can contribute significantly to nutrient uptake and direct observations confirmed that the crabs use this method in situ. The crabs are also able to use the setose appendages to sweep deposited material from the underlying substrate towards the mouth, presumably for nutritive purposes. The significance of these different feeding methods is discussed.     

Structure and function of the chelate pereiopods of the banana prawn Penaeus merguiensis

Penaeus merguiensis de Man systematically searches the substratum and removes small particles of food from it using the small chelae of the first three pairs of pereiopods. One or more chelae are used to transfer food particles to the mouth. The propus and dactylus of these limbs bear numerous setae arranged in discrete groups along the length of the segments. The tip of each seta is elaborately sculptured and has a large sub-terminal pore. It is probable that these are chemosensory organs responsible for discriminating edible from non-edible material. Proximal to almost every setal group is a much-branched seta (rarely two) which probably measures the depth to which the chela has been inserted into the substratum. At the articulation of the propus and carpus of the first pereiopod are three groups of differently sculptured setae, whose function is to clean the other chelate pereiopods. An elaborate system of pegs and a ridge on the apposable fringes of the chelae might be a mechano-receptive device with a particle size discriminating function. The ridges merge into opposable terminal denticulate pads whose function is to grasp food and other particles, which may be quite small (10 to 20 m upwards).     

Predaceous gastropods regulate new-shell supply to salt marsh hermit crabs

Marsh hermit crabsPagurus longicarpus Say directly acquire new shells as the predatory gastropodMelongena corona Gmelin consumes marsh periwinkles,Littorina irrorata Say. The influx rate of new shells into a salt marsh hermit crab population was measured by marking live periwinkles and daily recovering the shells from hermit crabs over periods of 3 to 6 d. Average rates of new shell acquisition ranged from 4.0 to 23.3 new shells per day from salt marsh areas of 4×10 m. Such consistently high rates contrast with the negligible rates generally assumed for new shell entry into hermit crab populations. The number of new shells acquired each day varied directly with the number of the predatory gastropod,M. corona, present in each study area at both natural and manipulated predator densities. Empty shells on the substrate are usually considered as the primary source of new shells to hermit crabs. However, over 500 empty shells had to be placed on the substrate in a 4×10 m area to provide a daily rate of 20 new shells to the hermit crab population.This is the first in a new contribution series from the Florida State University Marine Laboratory No. 1001     

How to bite the dust: morphology, motion pattern and function of the feeding appendages of the deposit-feeding thalassinid shrimp Callianassa subterranea

The morphology of the mouthparts of Callianassasubterranea (Montagu, 1808) was studied using light microscopy and scanning electron microscopy. All the mouthparts except the mandible, but including the mandibular palp, appeared to be supplied with a wide variety of setae. The setae of the medial rims of these appendages (the “ventral screens”) show a trend of decreasing passive motility towards the oral side; this applies to the setae themselves as well as the micro-structures on the setae. The motion of the mouthparts was filmed from aboral and rostral views with macro-video and endoscopy equipment, after marking the joints and tips of maxillipeds and the mandibles. Motion analysis showed that all mouthparts except the first maxilla (Mx1) actively moved during deposit feeding. The first and second maxillipeds (MP1, MP2), Mx2 and the mandibles moved at the same frequency, contra-lateral in phase, but with a 50% phase shift between ipsi-lateral mouthparts. The MP3 moved at a lower frequency with a contra-lateral phase shift of about 50% without an obvious phase relation with the other mouthparts. According to the following scenario, the feeding mechanism of C. subterranea seems to be based on the morphology and motion of the appendages in combination with specific setal functions. The MP3 make large excursions over the substrate, suspending the sediment in front of the MP2. The MP2 move through the upper layer of the suspension, trapping particles of 30 μm and smaller. These particles are combed out by the MP1 and transferred to the mouth opening by the cascade of finely structured setae of the inner mouthparts, and finally ingested. Received: 26 September 1997 / Accepted: 27 March 1998     

Obligate commensalism of <Emphasis Type="Italic">Curvemysella paula</Emphasis> (Bivalvia: Galeommatidae) with hermit crabs

Curvemysella paula is a markedly crescent-shaped bivalve that lives inside snail shells occupied by hermit crabs. Here, we describe the unique symbiotic life, growth pattern, and reproductive biology of this bivalve, based on specimens collected from the shallow, muddy bottom of the Seto Inland Sea, Japan. C. paula was found attached to columellae in the siphonal canal, mainly of nassariid snail shells occupied by two types of hermit crabs: Diogenes edwardsii (Diogenidae) and Spiropagurus spiriger (Paguridae). The crescent-shaped shell of C. paula is an adaptation to symbiotic life in the narrow interspace between the snail shell and the hermit-crab abdomen. C. paula is a protandric hermaphrodite. In our samples, each host snail shell harbored one (or rarely a few) large female and several males. All the female bivalves settled on the host shells with their anterior end facing outward and benefited from currents created by the hermit crab when feeding. In the muddy bottom, snail shells are a limited resource for both the hermit crabs and symbiotic bivalves. The bivalves benefit from the mobility of the hermit crabs, which prevent the shells from becoming buried in the mud. C. paula represents the only example of obligate commensalism with hermit crabs found in Bivalvia.     

Utilization of partially predated snail shells by the hermit crab Pagurus longicarpus Say, 1817

The hermit crab Pagurus longicarpus was shown to inhabit shells that were partially predated from intertidal areas of Long Island, New York. Among field collections of P. longicarpus, 2.13% of the hermit crabs (46 of 2155) were found with shells with snail tissue present. Over 90% of these partially predated snail shells were occupied by male hermit crabs. Although hermit crabs were in 8 species of snail shells, only Littorina littorea and Nassarius obsoletus were found occupied by hermit crabs and containing snail tissue. In the laboratory, we found that specimens of the spider crab Libinia emarginata were able to pull off the operculum of snails, leaving damage as found in field collections. In contrast, specimens of P. longicarpus were not able to prey on live, healthy snails. When specimens of P. longicarpus were placed in communal tanks, hermit crabs preferred partially predated snail shells to empty and original shells. However, original shells and empty shells were occupied with more frequency than partially predated shells when crabs were isolated. These findings indicate P. longicarpus actively seeks shells soon after attack and abandonment by snail predators, especially in the presence of competitors.     

Incidence,reproduction and feeding of Stylochus zebra,a polyclad turbellarian symbiont of hermit crabs

The hermit crab Pagurus pollicaris is the common host for the polyclad turbellarian Stylochus zebra in the Atlantic coastal waters from Massachusetts to North Carolina, USA. S. zebra is reported for the first time from two other hermit crabs, Pagurus impressus and Petrochirus diogenes. It was not found with Clibanarius vittatus or Pagurus longicarpus, although the latter serves as a host along the Gulf coast of Florida. The incidence of the polyclad with P. pollicaris in Massachusetts is more than twice that in populations to the south. Multiple infestations (up to 7 worms/crab) occur in over 50% of the infested crabs from Massachusetts, but they are rare elsewhere. Worms living in clear plastic shells with crabs orient in the body whorl dorsal to the host. Egg masses of worms are cemented in the same position within gastropod shells harboring the crabs. In the laboratory, the worm feeds on the gastropod Crepidula plana, a common inhabitant of shells occupied by P. pollicaris. Its feeding behavior is described. The relationship between S. zebra and P. pollicaris appears to be generally commensalistic, but it could be more complicated if the predation of crab embryos observed in the laboratory is confirmed under natural conditions.Contribution No. 755 from the Virginia Institute of Marine Science, Gloucester Point, Virginia, USA.     

Brown pelican siblicide and the prey-size hypothesis

Summary We asked whether the brown pelican (Pelecanus occidentalis) practices facultative brood reduction and tested two predictions of Mock's (1985) prey-size hypothesis: (1) if chicks take food directly from the parental mouth, nestmates should compete aggressively; (2) aggression between nestmates should increase during the developmental transition from indirect feeding (parents deposit food on the substrate) to direct feeding (parents pass food from mouth to mouth). Eggs in two-egg and three-egg clutches were laid and hatched 2 days apart. Junior (second- and third-hatched) chicks, fed less than their nestmates, grew more slowly and died more frequently. Of non-hatchling victims 74% were underweight, 55% bore peck wounds and 43% were found outside the nest, implying starvation, aggression and expulsion from the nest. Aggression occurred in all observed sibships, and junior chicks were subordinate in eight of ten cases. A progressive transition from indirect to direct feeding occurred during the 30 days after hatching, but there was no evidence of an increase in pecking during the transition. In conclusion, the brown pelican practices facultative brood reduction, and chick loss is mediated by sibling aggression as predicted by the preysize hypothesis. Pecking during indirect feeds seems to contradict the prey-size hypothesis, but may serve partly to establish a relationship of dominance-subordinance.Correspondence to: H. Drummond     

Food and feeding habits of the king crab Paralithodes camtschatica near Kodiak Island,Alaska

Stomach contents from 809 king crabs, Paralithodes camtschatica (Tilesius), from 6 areas near Kodiak Island, Alaska, and 9 sampling periods (1978–1979) were exammed quantitatively; 713 (88%) contained food. Mollusca (mainly the bivalves Nuculana spp., Nucula tenuis, and Macoma spp.) and Crustacea (mainly barnacles) were the dominant food groups in terms of percentage wet weight and frequency of occurrence; fishes were the next most important group of prey. No significant differences in feeding between sexes occurred; however, significant differences were apparent in the quantity of food consumed from different sampling periods, areas, depths, size groups, and crab molt-classes. Consumption was greater in spring and summer and in offshore locations at depths of 126 to 150 m. In addition, king crabs <140 mm carapace length (CL) consumed more food than crabs 140 mm CL. Adult, newshell (individuals that molted during the last molting period) females greater than 95 mm CL, and newshell males greater than 100 mm CL, each contained more food than did juvenile, newshell females <120 mm CL.Contribution No. 449, Institute of Marine Science, University of Alaska, Fairbanks, Alaska 99701, USA     

Shell-size selection by intertidal sympatric hermit crabs

This study evaluated selection for shell size by three species of tropical intertidal hermit crabs, Clibanarius antillensis, C. sclopetarius, and C. vittatus, from species of shells which are frequently used in nature. Crab size and weight were strongly and significantly related to all measured parameters of the selected shells. The strength of these relationships (r² values) depended neither on the crab nor on the shell variables taken into account. The relationships between crab size and the dimensions of the selected shells showed higher r² values than the corresponding relationships with the shells that the crabs had occupied when they were collected (0.482–0.903 in comparison to 0.091–0.652, respectively), indicating that the crabs were occupying sub-optimal shells in nature. Negative allometry was frequently found in the relationships between crab and shell variables, indicating that large crabs select and use proportionally lighter shells than do small crabs. This negative allometry was stronger for the shells used in nature (except for C. antillensis), i.e. larger crabs tended to select heavier shells in the laboratory than in nature. Different allometric relationships were also recorded among the dimensions of shells used in nature and those selected by the hermit crabs in free-access experiments: as shell length increased, the selected shells were heavier and had larger apertures than the shells used in nature. The relationships between crab size and the length and weight of the selected shells did not depend on the species of crab or species of shell, but only on crab size. Therefore, analyses using these variables can be performed without taking the species of crab or shell into account, i.e. data from different crab or shell species can be pooled. The influence of crab and/or shell species was recorded only in the models fitted for aperture length and width, variables which were more related to shell architecture than did shell length or weight. In contrast, if crab weight is used as an independent variable, different crab or shell species can be analyzed together independently of the particular shell parameter. This indicates that crab weight may be less susceptible than crab shield length to shell morphological constraints. Finally, the results indicate that the preferred size of a given shell type chosen by a given hermit crab will depend more on crab size or weight, than on the crab or shell species under consideration, i.e. crab shell-size relationships are not species specific.Communicated by P.W. Sammarco, Chauvin     

Shell acquisition by hermit crabs: which tactic is more efficient?

Gastropod shells are limited resources for most hermit crab species, acting as primary factors in various aspects of their biology. To investigate the efficacy of different behavioral tactics adopted for their acquisition (locomotion, attendance at shell-supplying sites, interactions with conspecifics in aggregation) we conducted observations and experiments at a salt marsh in New England (USA). Locomotion, fast and meandering, significantly increased the chances of encountering empty shells and conspecifics. However, shell exchanges were rare. Simulated gastropod predation sites quickly attracted a larger number of hermit crabs than the other shell-supplying sites tested (live and dead conspecifics, live snails) and induced the rapid occupancy of all the empty shells offered, usually by the first crabs arriving at the site. Aggregations seemed not to function as shell exchange markets, as previously suggested for several other species. In the short run, exploitation seems to be more efficient for the acquisition of new shells by Pagurus longicarpus. In the long run, it is the density of nondestructive gastropod predators that regulates the availability of new shells of good quality in the pool available to this hermit crab species.     

Locomotion,feeding, grooming and the behavioural responses to gravity,light and hydrostatic pressure in the stage I zoea larvae of Ebalia tuberosa (Crustacea: Decapoda: Leucosiidae)

The stage I zoeae of Ebalia tuberosa swam by sculling with the exopodites of the 1st and 2nd maxillipeds and flexed the abdomen to brake or change direction. The larvae gained depth by stopping all natatory movements and sinking passively at rates of 6 mm s^-1. The zoeae refused both living and dead nauplii of Artemia spp., as well as two species of diatoms, but fed readily on detritic material on the bottom which they scooped up using the endopodites of the maxillipeds and pressed against the mouthparts using the telson. The setae on the posterior border of the telson were used for grooming the maxillipeds and the anterior mouthparts. Day-old stage I zoeae were negatively geotactic, positively phototactic and responded to pressure increases by swimming upwards and by high barokinesis. By the third day some larvae had become positively geotactic but were photopositive, and the majority responded to pressure increases as in the day-old larvae. Five-day old larvae were still photopositive but the majority had become positively geotactic and fewer himbers responded to pressure. Seven-day old larvae failed to respond to any of the stimuli used and assumed a predominantly benthic lifestyle. It is suggested that this anomalous behaviour is related to the dispersal of the larvae and to the specialized habitat requirements of the adults while the rather unusual morphology of the larvae is related to their feeding behaviour and semi-benthic lifestyle.     

Changes in the feeding behavior of the deposit-feeding polychaete<Emphasis Type="Italic"> Laeonereis acuta</Emphasis> on soft sediments inhabited by burrowing crabs

In this paper, we evaluate the influence of the burrowing crab Chasmagnathus granulatus on the feeding behavior, body condition, burrowing activity, and fecal production of the deposit-feeding polychaete Laeonereis acuta. Previous results and ours show that, due to crab activity, sediment organic matter decreases with depth outside a crab bed, but there were no differences inside. Also, particle sizes were smaller inside the crab bed. Polychaetes showed better body condition inside the crab bed, suggesting higher energy gain. They fed on the surface more frequently outside the crab bed than inside. However, feeding rate was higher inside the crab bed, which is consistent with the larger weight and volume of fecal pellets found inside crab bed. In both areas, the bedload sediment transport was not similar to the sediment ingested by this polychaete. A 1-month exclusion/inclusion experiment performed outside and inside the crab bed showed that the body condition of polychaetes changed between areas but that this was not directly due to crab manipulation. Thus, our results suggest that the higher sediment quality inside a crab bed can positively affect the feeding behavior of the deposit-feeding polychaete L. acuta, increasing its feeding rate and consequently enhancing its body condition.Communicated by P.W. Sammarco, Chauvin     

Feeding factors for the sea anemone Anthopleura midorii

The carnivorous feeding behavior of the sea anemone Anthopleura midorii was found to be a sequence of successive feeding motions, which consisted of several separate actions: (1) tentaculation on any solid matter; (2) retention of prey by tentacles; (3) mouth opening; (4) ingestion of food; (5) digestion of food; (6) extrusion of indigestible waste material. Hot water extract of a natural prey induced the entire feeding sequence when extract was given in the form of an artificial food by mixing with -potato starch and water. The fractionated extract induced only a limited feeding action. Several amino acids induced feeding. Alanine, glycine or histidine stimulated the retention of artificial food by tentacles, and proline evoked mouth opening. Cysteine or reduced glutathione (GSH) induced food ingestion. Unnatural food stuff such as -potato starch was ingestible, if an appropriate amount of a chemical substance was added.     

Inhibition of marine algal photosynthesis by heavy metals

The orientation of the opportunist, sublittoral barnacle Balanus trigonus is investigated. Water movement is the primary orientation stimulus for this species, but this effect may be modified by the influence of light acting in opposition to it. The barnacles orientate at right angles to the axis of wave-surge movement, and observations of feeding behaviour showed that in this position the cirral net could be swivelled 90° each way to make best use of both the advancing, and the reversing, water flow. In a steady unidirectional tidal current the barnacles aligned themselves so as to beat with the water flow, and observation suggests that stroking with the flow is more efficient than forcing the cirral net against the motion of the water. It is suggested that, for an opportunist short-lived species such as B. trigonus which is subjected to high predation rates, there is considerable selection pressure for rapid growth and early sexual maturity. To achieve this in a filterfeeding organism for which food is limiting, food collection must be as efficient as possible. B. trigonus has adopted a system of orientation to a variety of water movement regimes that allows efficient food collection over a wide range of conditions.     

Ecology of subtropical hermit crabs in SW Madagascar: short-range migrations

Many mobile animals migrate because of the different benefits provided by different localities in time and space. For hermit crabs, such benefits include resource (shell, water, food) acquisition and gamete release. One of the more successful crustacean land-invaders, Coenobita hermit crabs, undertake complex short-range migrations in SW Madagascar. Number of active hermit crabs was inversely related to wind strength and positively related to tidal range, emphasising that movement would conserve water. A circadian component was also recorded in the locomotory activity of Coenobita pseudorugosus and C. rugosus. Path linearity varied with many of the same parameters, but also with beach slope. Movement was primarily perpendicular to shore in small individuals, but the parallel proportion increased with hermit crab size and tidal range, probably driven mostly by shell and food searching. Despite the costs of movement and shell carriage in the terrestrial environment, C. pseudorugosus and C. rugosus were as fast as their marine counterparts. Their speeds varied principally with individual size and were approximately 20% faster without shells and about 20% slower when climbing up a 20° slope, compared to horizontal or downhill travel. Hermit crabs, which are highly numerous and speciose in SW Madagascar, do not seem to partition niches by differential movement patterns. Aside from provision of shells in middens and capturing large adults for bait or pets, human activity may have a profound effect on hermit crab movement: observations at rare uninhabited marine reserves like Nosy Ve show that considerable diurnal activity may take place despite the apparent hostility of the environment to an essentially marine animal.     

Orientation of Coenobita rugosus (Crustacea: Anomura): A field study on Aldabra

Experiments were designed to determine whether anemotaxis shown by the terrestrial hermit crab Coenobita rugosus H. Milne Edwards is (i) an orientational strategy leading the crabs to sea or (ii) a strategy improving rectilinear flight. In an arena with uniform landscape, hermit crabs from different beaches of the Aldabra atoll orientated during daylight in the direction of the sea of their own beach; this orientation was only slightly affected by wind direction. Under artificial wind conditions, at night, all hermit crabs displayed positive anemotaxis, independent of the orientation of the home beach, thus leaving little support for the first hypothesis. During the day, with the sea and beach both visible, all the hermit crabs on the beach moved landwards irrespective of wind and home beach direction.     

Mechanosensory properties of the mouthpart setae of the European shore crab <Emphasis Type="Italic">Carcinus maenas</Emphasis>

In decapod crustaceans, the largest density and diversity of sensilla, referred to as setae, is in general found on the mouthparts, but little is known about their sensory properties and thereby their functions. Here data are presented from mechanoreceptors from the two largest mouthparts, maxilliped 2 and 3, of the European shore crab Carcinus maenas. The mechanoreceptors were found to respond to either displacements of the entire seta or bending of the setal shaft. The displacement-sensitive cells encode both the amplitude and the velocity of the displacement and about half were found to be directional but most in a non-exclusive way. The amplitude of the stimulus is encoded in the number of spikes produced with a linear correlation. The velocity is encoded in the interspike intervals with shorter intervals at higher velocities. In the latter case, the correlation follows a power function. The physiological data is correlated with the morphology and usage of the maxillipeds were examined with scanning electron microscopy and macro video recordings respectively. Recordings were obtained from cells associated with four different setal types and they all showed similar mechanosensory properties supporting that the external morphology of setae is more closely connected to their non-sensory functions, e.g., mechanical manipulation of the food items. The details of the sensory properties together with the high setal density, especially on maxilliped 3, suggest that a large amount of tactile information is gathered during feeding.     

Epibiosis on cerith shells in a seagrass bed: correlation of shell occupant with epizoite distribution and abundance

The cerith Cerithium atratum (Born 1778) is an abundant gastropod in the seagrass beds at Cabo Frio, Brazil. In order to estimate the ecological importance of cerith shells as a rare hard substratum in the seagrass bed, the abundances of C. atratum and of cerith shells occupied by hermit crabs were quantified. The mean densities of C. atratum and hermit crabs were 1887 and 100 individuals m⁻², respectively, and these provided 0.5 m² shell area m⁻² available for epizoite colonization. The tube-forming polychaete Hydroides plateni (Kinberg 1867) and oyster Ostrea puelchana Orbigny, 1841 were the dominant visible epizoites on inhabited cerith shells. These epizoite populations were compared in order to investigate whether the temporal and spatial patterns in the epibiotic community were related to ecological and behavioral aspects of the occupant species (cerith or hermit crab). Larger cerith shells had a greater abundance of epizoites. Each epizoite showed a preference for a different occupant of the shells (the oyster for C. atratum and the polychaete for cerith shells occupied by hermit crabs). The oyster showed a seasonal pattern in abundance on C. atratum, being more common in fall (March–April). The distribution of the epizoites on the shells depended on the shell occupant species and was probably related to their different foraging activity –C. atratum ploughs half buried through the sediment, while the hermit crab crawls on the sediment surface. In both cases, the activity of the shell occupant was considered to be beneficial to the epizoites, as empty shells and shell fragments did not support a macroepifauna. Received: 1 May 2000 / Accepted: 8 August 2000