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).     

Settlement behavior and antennulary biomechanics in cypris larvae of Balanus amphitrite (Crustacea: Thecostraca: Cirripedia)

The settlement behavior, antennulary biomechanics and morphology were investigated in the intertidal cypris larvae Balanus amphitrite. The very maneuverable antennule of the cyprid consists of four jointed segments, and it is the prime appendage for substratum exploration and final settlement. We recorded several previously undescribed settling behaviors, such as the "tight direction change", for this intertidal species and discovered that the antennulary morphology is modified to facilitate such behaviors. Segment 2 has a modified lateral cuticle, which gives the segment an additional higher degree of maneuverability compared to non-intertidal cirripede species of Rhizocephala. Additional supporting muscle groups were also found in the antennule of our species, which were not found in the investigated rhizocephalan larvae. The results could very well be associated with the higher needs of substratum recognition in intertidal species, due to more heterogeneous habitats such as intertidal rock flats.     

Relationship between mouthpart structures and in situ feeding habits of species of the family Pontellidae (Copepoda: Calanoida)

Pontellid copepods were collected from the surface waters of a tidal front region in the Bungo Channel (the Inland Sea of Japan) in June 1986 to examine the relationship between the morphology of cephalic appendages and gut contents. In particular, two dominant species,Labidocera japonica Mori andPontellopsis yamadae Mori, were compared in detail. Large setae on the second maxillae ofL. japonica possessed two rows of setules at right angles to each seta along its inner margin except a terminal part which was serrated, whereas the inner margin of those setae ofP. yamadae was entirely serrated. Judging from the structure of the mouthparts, especially the second maxillae, the former species seems to employ both suspension and raptorial feeding modes, in contrast to the latter, which may use only the raptorial mode of feeding. InP. yamadae, the first maxilla and the maxilliped are also modified for carnivory. Gut content analysis supported the morphological evidence for feeding differences, and revealed thatP. yamadae is a carnivore preying mainly on copepodids whileL. japonica feeds omnivorously on copepod nauplii and phytoplankton particles. Since the mouthpart structures of congeners are quite similar to each other, the feeding behavior and habits might also be similar. Within the family Pontellidae, the generaAnomalocera, Calanopia, Epilabidocera, andPontella have mouthpart structures similar to those ofLabidocera, whereas the genusPontellina resemblesPontellopsis. Morphological similarities would suggest that the first group of genera employs both suspension and raptorial feeding modes, and thatPontellina is a carnivore likePontellopsis.     

Influence of cultivation and food concentration on body length of calanoid copepods

The cephalothorax length of the marine pelagic Copepoda Acartia clausi Giesbrecht, Temora longicornis (Müller), Centropages hamatus (Lilljeborg) and Pseudocalanus sp. was monitored at 15°C during prolonged cultivation through up to 55 filial generations and at different concentrations of food. The length of T. longicornis decreased considerably during the first 15 generations and remained rather constant thereafter. In the other species, body length increased slightly or remained almost constant. Genetic changes are probably involved. Food concentration influenced body size of all species, particularly C. hamatus, in which 80% of the natural size range may be explained by differences in food concentration. The idea that temperature is a dominant factor in determining the length of copepods should be reconsidered.     

Regulation of body dimethylsulfoniopropionate (DMSP) content by the copepod Temora longicornis: a test of four mechanisms

Copepods contain dimethylsulfoniopropionate (DMSP) in their tissues in addition to the DMSP in their gut contents and therefore constitute an additional pool of DMSP in the ocean. In the estuarine copepod Temora longicornis Müller, this dynamic pool of DMSP is reduced when external salinity decreases. In the present study the mechanism(s) used by T. longicornis to adjust its DMSP content were examined. Four possible mechanisms were tested in experiments conducted between July 1997 and May 1999: (1) DMSP cleavage, (2) demethylation, (3) conversion to methionine, and (4) release from the body. Tissue extracts of T. longicornis did not exhibit the ability to cleave or demethylate DMSP. In incubation experiments, when external salinity decreased, T. longicornis individuals reduced their DMSP content without increasing their methionine content. The total amount of DMSP in the incubation vials was conserved regardless of salinity. The copepods retained most of the DMSP in their tissues in 30‰ water, but when salinity was reduced to 20‰, the copepods released 41 to 55% of their body DMSP to the surrounding medium; 89 to 91% of the total DMSP was recovered. This suggests that estuarine copepods represent a reservoir of DMSP, which is released as dissolved DMSP upon salinity decline. Therefore, while osmoregulation by these copepods may have little impact on the chemical properties of DMSP, it will affect the partitioning of DMSP between particulate and dissolved phases in the water column. Received: 20 August 1999 / Accepted: 28 March 2000     

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.     

Antifouling adaptations of caridean shrimps: Cleaning of the antennal flagellum and general body grooming

Structural specializations of chelipeds used by caridean shrimp in two kinds of grooming activity are described. In general body grooming, the chelipeds, and in some species, the last walking legs, nip, pick or brush material from the exoskeleton. When the cleaning chelipeds are the second pair, the carpal segment is multisegmented, increasing distal flexibility which aids in grooming. Tufts of compound setae and setal chela locks are characteristic of cleaning chelae. In representatives from 13 of 15 caridean families surveyed, brushes of serrate setae surround the carpal-propodal joint of the first cheliped. This setal structure is used in the specific task of cleaning the chemotactile antennal flagellum. Cleaning brushes on the last walking legs of some species are involved in general body cleaning. Experiments on Heptacarpus pictus showed that when the cleaning chelipeds were ablated, body parts became fouled with epizoites and particulate debris in experimental shrimps, while control shrimps showed little fouling. Suggestions on the adaptive role of general body cleaning in these natant animals are discussed. A survey of cleaning characters in representatives from 15 caridean families suggest that such characters are rather constant within a family. A possible correlation between the taxonomic success of a family and the degree of development of general body cleaning is suggested.     

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.     

Feeding ecology of Liocarcinus depurator (Decapoda: Portunidae) in the Ria de Arousa (Galicia,north-west Spain): effects of habitat,season and life history

The diet of the portunid crab Liocarcinus depurator (L.) in soft-bottom areas of the Ría de Arousa (Galicia, north-west Spain) was studied by analysis of stomach contents, and by comparison of habitats of contrasting abundance, composition and diversity of prey communities. Monthly samples were taken from July 1989 to June 1990 (N=3747) in mussel (Mytilus galloprovincialis) raft-culture areas and central channel zones. Ninety-two food items were identified: crustaceans, molluscs, polychaetes, ophiuroids and fishes were the dominant prey. The raft epifauna and the mussels constituted the dominant group of prey in the raft areas and inner channel station (the epifaunal crab Pisidia longicornis comprised 30 to 52% of the diet), whereas the benthic macroinfauna was the dominant food component in the mid-outer channel (polychaetes constituted 32% of the diet). In the epifaunal community, diet selection for P. longicornis was positive and for amphipods was negative, whereas within the infauna diet selection for bivalves and ophiuroids was positive and for polychaetes negative or neutral. The seasonal importance of P. longicornis and M. galloprovincialis varied in the diet of L. depurator parallel to seasonal activities connected with mussel culture (seeding, thinning, harvesting) that determine the availability of prey from the rafts. Major changes in the composition of the diet related to the life history of L. depurator occurred during ontogeny. In the raft zones, consumption of mussels and fishes increased with increasing body size, while the consumption of Natantia, non-decapod crustaceans, gastropods, polychaetes Pectinariidae and plants decreased with increasing crab size. Maximum consumption of P. longicornis was by L. depurator individuals of intermediate size (15 to 44 mm carapace width). Stomach fullness (measured as dry weight of food) increased with increasing body size, but with negative allometry. During the late pre-moult and early post-moult periods L. depurator did not consume any food; maximum stomach fullness was recorded for the late post-moult and intermoult stages. Female food consumption decreased during egg incubation. Food consumption was greater in autumn and winter than at other seasons; this appears to be related to seasonal changes in reproduction and moulting.     

Fine structure of the “button setae” in the deep-sea pelagic copepods of the genus Euaugaptilus (Calanoida: Augaptilidae)

 Many species of the carnivorous copepod family Augaptilidae, a primarily meso- and bathypelagic group, possess specialized “buttons” on the setae of their maxilla and maxilliped, which have been thought to be a simple cuticular outgrowth. The fine structure of the button setae in three species of Euaugaptilus was examined by light- and electron microscopy from samples collected in the eastern Indian Ocean and the subtropical western North Pacific. The buttons are arranged in two rows along the inner surface of the setae. There are differences in the shape, size, and arrangement of buttons among the three species, Euaugaptiluslaticeps, E.longimanus, and E.magnus. The button setae have an elaborate internal structure consisting of “setal shaft”, “stalk”, “disc”, and “outer membrane”, with microtubule bundles in the setal lumen, but have no muscular tissues. The disc has only pin-point contact to the core of the stalk, but has connection over a wider area with the fibrous tissue surrounding the core, and the contact area between the stalk and the setal shaft is also small. These structures suggest that the disc will move freely on the apex of the core of the stalk and the stalk may be movable at its connection to the setal shaft. All these components of the button setae may function in concert to fit the surface of the discs firmly to the surface of a prey, and to absorb the energy generated by the struggling prey. Received: 30 November 1999 / Accepted: 19 May 2000     

Overwintering strategies of dominant calanoid copepods in the German Bight, southern North Sea

Abundance, stage composition and reproductive parameters (egg production, egg viability, proportion of spawning females) of the four copepod species Acartia clausi, Centropages hamatus, C. typicus and Temora longicornis were measured at the long term sampling station Helgoland Roads (German Bight, southern North Sea) from September 2003 to May 2004 to study their overwintering strategies. A. clausi was overwintering as females with arrested reproduction from November to January. T. longicornis, which is known to produce resting eggs in the North Sea, had a pelagic population with all developmental stages present during winter and reproductive rates closely related to food concentrations. Although their females produced eggs in response to ambient food conditions, both C. hamatus and C. typicus were rare in the pelagic. The C. hamatus population returned in May, probably from resting eggs, whereas C. typicus depended on advection. The Centropages species seemed to be less adapted to pelagic life in winter than A. clausi and T. longicornis. Sporadic occurrence of large numbers of nauplii and young copepodids of A. clausi and Centropages spp. pointed to different overwintering strategies or more successful survival in adjacent regions and advection of them into the waters around Helgoland island. While A. clausi was decoupled from environmental conditions in late autumn and winter, the other species were able to respond to variations in the food environment. Thus, egg production of T. longicornis increased during an unusual autumn diatom bloom.     

Population structure and breeding biology of Pisidia longicornis and Porcellana platycheles

Quantitative sampling of littoral and sublittoral populations of Pisidia longicornis (L.) and a littoral population of Porcellana platycheles (Pennant) has been carried out over a 2 year period (1970/1972). On the shore, P. longicornis were most abundant at mean low water spring (MLWS), whereas P. platycheles were found mostly at lower mid-shore level. Differences were found between littoral and sublittoral P. longicornis. Sublittoral females are gravid for a slightly shorter period (April to September), carry more eggs, and are smaller than littoral females. It is suggested that their lifespan may also be shorter than that of littoral individuals. Young of both species settle in autumn and grow until December, when growth ceases until the following spring. Numbers decline in the summer months due to the mortality of older individuals. Many sublittoral and some littoral P. longicornis are sexually mature towards the end of their first year on the shore, and may copulate after the early spring moult and carry eggs in the subsequent summer months. P. platycheles appear to follow a similar basic pattern, but results are less clear.     

Semiochemicals mediating host-finding behaviour in the phoretic association between Macrocheles saceri (Acari: Mesostigmata) and Scarabaeus species (Coleoptera: Scarabaeidae)

Summary. Phoresy is an interspecific association where mobile organisms ensure small and/or wingless organisms to colonize patchy distributed environment. Macrocheles saceri (Acari: Mesostigmata) is a phoretic mite which specialized on Scarabaeus dung beetles (Coleoptera: Scarabaeidae). Variation for phoretic load and sites for mite fixation on beetle body has been tested according to Scarabaeus species. The number of setae on beetle body cannot explain the observed patterns in mite fixation. Evidence was provided that the semiochemicals of the host cuticle play a kairomonal role in the host finding behaviour of phoretic mites. In addition semiochemicals ensure host discrimination.     

Differences in reproductive strategies of morphs of the genus Echinaster (Echinodermata: Asteroidea) from the Eastern Gulf of Mexico

Two morphs of the asteroid genus Echinaster, E. Type 1 and E. Type 2, have reciprocal annual cycles of reproduction and nutrient storage. The pyloric caeca (storage organs) reach maximal size over winter then decrease with increasing gametogenic activity in the spring. E. Type 1 broadcasts buoyant eggs in late May to early June. E. Type 2 deposits benthic eggs in late April. Lecithotrophic larval development is similar in both morphs except that larvae of E. Type 1 undergo a brief planktonic phase. E. Type 1 reaches sexual maturity at a larger size, expends a lower reproductive effort but has a greater absolute reproductive output than E. Type 2. E. Type 2 produces fewer, larger eggs and has a greater parental investment per egg. In both morphs, females have much higher gonadal lipid levels than males and expend a higher reproductive effort in terms organic matter and energy. Reproductive effort, reproductive output, nutrient storage in the pyloric caeca and body size varied between populations and between years in each morph. Egg size and parental investment per egg were constant. Within-morph variability is attributed to differences in nutritional state. Differences in reproductive strategy support the hypothesis that the morphs are separate species.     

Comparative morphology and cytology of siphons and siphonal sensory organs in selected bivalve molluscs

 The form, anatomy and cytology of the water-passages in siphon-possessing and siphon-lacking species of a selected group of bivalve molluscs from the Red Sea (Callista florida, Circe currogata, C. crocea, Lucinia dentifera, Dosinia histrio, Pitar hebraea, Tridacna maxima, Pteria aegyptica), and from the Mediterranean Sea (Spondylus spinosus, Pinctada radiata, Pinna nobilis, Donax trunculus, D. semistriatus, Mactra stultorum, Tapes decussatus, Petricola lithophaga, Brachidontes pharaonis) were studied by means of light and electron microscopy. In the mytiloids, ostreoids and ptrioids studied, the water-passages are covered by a ciliated epithelium and the few tentacles around their openings are identical to those found on the mantle edges. Contrary to this, in the veneroids studied, the siphons are covered by a microvillar epithelium and their tentacles, especially of the inhalant siphon, reveal characteristic branching. The so-called Type I, Type II and Type III ciliated sensory organs on and within the siphons are similar to those observed in other bivalves, differing from each other in the dimension, number and form of cilia. In the studied mytiloid species only Type I and Type II organs were found. The form and structure of the siphons and the distribution of sensors on them can be used as markers for ecological as well as phylogenetic considerations. Received: 28 January 2000 / Accepted: 28 May 2000     

Feeding and mouthpart morphology in the semaphore crabHeloecius cordiformis (Decapoda: Brachyura: Ocypodidae)

The mouthparts of the Australian semaphore crabHeloecius cordiformis (H. Milne-Edwards, 1837) are described. Male and female semaphore crabs were observed and collected from mangrove swamps around the south side of Botany Bay, Australia between 1983 and 1987. Semaphore crabs inhabit mangrove mud/sand flats between the midlittoral intertidal zone and the supralittoral fringe of bordering saltflats. Scanning electron micrographs provide detailed pictures of mouthpart morphology and the setae present on the third, second, and first maxillipeds; the second and first maxillae; and the mandibles. Amongst other setal types, spoon-tipped, corn, and bottle-brush setae are described. While semaphore crabs are primarily deposit feeders (microphagous), they have well developed mandibles and are, therefore, also able to eat larger plant and animal material (macrophagous). A brief comparison is made between the mouthparts of semaphore crabs and fiddler crabs (Uca spp.), since the two genera are closely related, occupy similar niches, and resemble each other in appearance. This comparison reveals that while the second maxilliped setal morphology differs between the two genera (Heloecius possesses corn setae,Uca possesses spoontipped setae) these setae may have similar functions. The feeding behaviour ofH. cordiformis is related to substrate composition, burrow construction and various mouthpart specialisations. It is conlcuded that, within the context of the present study, certain structural e.g. mouthpart modifications allowing both scavenging and deposit feeding, and behavioural modifications e.g. mound building in wet muddy habitats, enable semaphore crabs to occupy a wide range of habitat types within the mangrove.Please address all correspondence and requests for reprints to Dr Maitland at his present address at the University of the Witwatersrand     

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     

Complex chemical communication in the crazy ant <Emphasis Type="Italic">Paratrechina longicornis</Emphasis> Latreille (Hymenoptera: Formicidae)

Summary. The formicine ant Paratrechina longicornis is known for its extremely opportunistic foraging behaviour. Only a single trail pheromone source, the rectum, was previously described from this ant. Our detailed examination of this ant’s chemical communication system revealed the presence of at least four sources of pheromones. Rectum, poison sac, and Dufour gland contain orientation components with decreasing effectiveness and persistence (in the sequence mentioned) as well as attractants with increasing effectiveness. Furthermore, the mandibular gland contains repellents, and a releaser of defensive behaviour. This set of various signals of different strength and persistence allows an extraordinary degree of flexibility and efficiency in the collective behaviour of P. longicornis, especially food exploitation, and thus may contribute to this insect’s overall ecological success.     

Distribution of overwintering calanoid copepod eggs in sea-bed sediments around southern Britain

Eggs of the calanoid copepodsLabidocera wollastoni (Lubbock) andCentropages hamatus (Lilljeborg) and others, which could not be reliably assigned to species, were identified from sediment samples taken during three cruises in December 1984, January 1986, and February 1987 in the southern North Sea, the English Channel, the Irish Sea and the Celtic Sea. Eggs were most abundant in sediment samples taken between 20 and 80 m depth, with bottom stress less than 10 dyn cm^–2, and near to the positions of tidal fronts. It is suggested that sedimentation, resuspension and sediment transport contribute to the causes of these distributions, which have been found to be consistent with the distributions of species which are known to produce diapause eggs as an overwintering strategy. Nauplii ofL. wollastoni, C. hamatus, C. typicus (Krøyer)Temora longicornis (Müller) andAcartia sp. hatched from incubated sediment samples. The last noted hatchings ofL. wollastoni, Centropages spp. andT. longicornis all occurred more than 1 yr after the start of the incubation experiments.