Ciliary feeding of tornaria larvae of Ptychodera flava (Hemichordata: Enteropneusta)

Ciliary feeding by tornariae of Ptychodera flava (Eschholtz) and other tornariae from the plankton is compared with Garstang's (1939) account of feeding by these larvae, which account contains errors, and with ciliary feeding by echinoderm larvae. Some details of ciliation are also described. As in echinoderm larvae, band cilia beat away from the food grooves and retain particles on the upstream side of the ciliated band, but tornariae use muscles less in ingestion and rejection. In early-stage and most late-stage echinoderm larvae, the ciliated band functions in both swimming and feeding, but in tornariae the ciliated band is arranged meridionally so that few portions of the ciliated band produce a posteriorly-directed current and a locomotory teletroch is needed for swimming. Faster-swimming tornariae observed in bowls achieved higher ingestion and clearance rates. These observations raise questions about form and function in the giant Planctosphaera pelagica. Cilia of the locomotory telotroch increase in length as tornariae increase in size, but there is little increase in length of cilia which capture food. Instead, the length of the ciliated band increases relative to other larval tissues by means of increasing convolution of the band. Hence, the volume of water processed for food probably increases relative to energy expended by larvae during development to the tentaculate stage. However, the length of the ciliated band may decrease relative to other larval surfaces with continued increase in size beyond this stage. These interpretations of growth and feeding efficiency are consistent with the reported geographic distribution of tornariae with and without tentacles.     

Function of lateral cilia in suspension feeding of lophophorates (Brachiopoda,Phoronida, Ectoprocta)

Suspension feeding by the brachiopod Laqueus californianus (Koch), the phoronid Phoronis vancouverensis Pixell, the ectoprocts Bugula sp., Membranipora villosa Hincks, and Schizoporella unicornis (Johnston), and actinotrocha and cyphonautes larvae was observed. Lophophorates retain particles on the upstream side of the band of lateral cilia, even after particles have moved lateral to the frontal surface. This suggests that these animals utilize an induced local reversal of beat of the lateral cilia for concentration and capture of suspended food particles. Retention of particles can cease while the current past the tentacles continues. Movement of particles down the frontal surface of the tentacles of L. californianus and the ectoproct species confirms previous observations that mucus strands are not used in particle capture. Possible functions of latero-frontal cilia or bristles are considered. Distribution of the feeding mechanism among phyla, clearance rates, and the lack of fusion of tentacles in brachiopods are discussed. The impingement mechanism previously suggested for lophophorates cannot account for the movements of particles observed here.     

Role of particle wettability in capture by a suspension-feeding crab (Emerita talpoida)

Suspension feeders sometimes depend on adhesion between particle and collector to capture food. If food particles have different adhesive properties than other particles, food could be passively selected by adhesive mechanisms. In this regard, the effect of particle wettability on adhesion to artificial and natural collectors was studied. First, the adhesion of glass particles to artificial collectors, both varying in wettability, was assessed to determine if wettability influenced adhesion in seawater. The adhesive force between glass particles and artificial collectors was measured by increasing the force pulling particles away from the collector until 50% of the particles fell off the collector. Adhesion increased as particle wettability decreased. Next, glass particles were used to determine if the antennal collector of the suspension-feeding mole crab Emerita talpoida captured particles based on particle wettability. 0.5 to 10 and 15 to 25 μm particles were suspended in a recirculating flow tank filled with seawater, and ablated antennae were exposed to this flow, after which the captured particles were counted. Results for the 0.5 to 10 μm particles confirmed predictions based on results from artificial collectors; particle capture increased as particle wettability decreased. The 15 to 25 μm particles may have been captured by sieving, and consequently did not follow predictions based on adhesion. Passive selection of particles based upon wettability differences can occur. Received: 17 April 1998 / Accepted: 14 September 1998     

Gill function and particle transport in Placopecten magellanicus (Mollusca: Bivalvia) as revealed using video endoscopy

The technique of endoscopic video observation was used to study feeding processes of Placopecten magellanicus (Gmelin), collected from Bull Arm, Newfoundland in August 1991 and 1992, under near-natural feeding conditions. The fate of captured particles depended on the extent of ingestive or handling capacity saturation. Under low (1 to 10 particles l^-1) to medium (10 to 20 particles l^-1) particle concentrations, most particles were incorporated in continuous anteriorly directed slurries in the dorsal ciliated tracts of the gill arch and dorsal bends. As particle concentration or exposure time to the lower particle concentrations increased, four endogenous mechanisms of ingestion volume control were increasingly observed: (1) rejection of dense mucus-particle masses from the principal filament troughs onto the ventrally beating cilia and associated currents of the ordinary filament plicae, counter to and below the incoming pallial current maintained by the principal filament cilia; (2) intermittent stopping of the anteriorward flow in the dorsal ciliated tracts; (3) reduction or cessation of input from the principal filaments to the dorsal ciliated tracts; (4) detachment of the dorsal bends from the mantle to establish a shunt from the infrabranchial to the suprabranchial cavity. Chemical and histochemical tests of purified fluid withdrawn from the dorsal ciliated tracts indicate that mucus is present at all particle concentrations. Mucus therefore participates both in normal feeding and in ingestion volume control on the bivalve gill, although different mechanisms, and types of mucus, effect transport of material in the dorsal (feeding) and ventral (cleaning) ciliated tracts.     

Reduced planktotrophy in larvae of <Emphasis Type="Italic">Clypeaster</Emphasis><Emphasis Type="Italic">rosaceus</Emphasis> (Echinodermata,Echiniodea)

Phylogenetic analyses have demonstrated that nonfeeding larvae have evolved from feeding larvae many times among marine invertebrates. In light of this observation, it is surprising that an intermediate strategy, a larva that can feed but is provisioned with enough energy to metamorphose without acquiring exogenous food (i.e., facultative planktotrophy), is rare. A hypothesis for the lack of facultative planktotrophic species among marine invertebrates is that the transition from feeding to nonfeeding is rapid due to this intermediate stage being evolutionarily unstable. Evidence that would support this hypothesis is if species with facultative planktotrophy have reduced food assimilation when compared with obligate planktotrophs. We studied a species with facultative planktotrophic larvae, Clypeaster rosaceus, that is very near the boundary between facultative and obligatory planktotrophy, to answer two questions: (1) does feeding during the larval stage result in energy gains in larval or juvenile stages and (2) if not, are larvae capable of assimilating exogenous food at all. Our measurements of energetics in larval and juvenile stages show that C. rosaceus larvae accumulate very little if any energy when fed, but stable isotope data indicate that larvae are able to assimilate some food. Our results are consistent with similar studies on facultative planktotrophic larvae suggesting poor food assimilation and rapid loss of larval feeding after a population evolves the ability to reach metamorphosis without feeding (lecithotrophy).     

Functional morphology of the podia and ambulacral grooves of the comatulid crinoid Antedon bifida (Echinodermata)

Feeding units — viz. triplets of unequally-sized podia associated with protective lappets — occur all along the pinnules of adult Antedon bifida (Pennant). Small food particles are trapped by direct mucus impingement to the wall of primary and secondary podia (there are no mucus net or mucus thread helping in this process). Large particles are caught by primary and secondary podia which partly curl over them. Small particles accumulate on the collecting podia before being transferred to the groove, while large particles are transferred one by one. Transfer of particles to the groove occurs by wiping the collecting podia on the ciliary tracts against the ciliary current. When active, tertiary podia always paddle against the ciliary current. They serve in bolus formation (mucus embedding of food particles), but do not participate in bolus compaction or propulsion. Elimination of unwanted particles occur through the action of secondary podia whose movements may disrupt the lappet's palisade, thus creating a sideward current that moves particles from the groove to the outside. Typical podial triplets do not occur along brachial and calycinal grooves. Both brachial and calycinal podia function mostly in guiding and regulating particle flow. The feeding structures of early pentacrinoid larvae of Antedon bifida recall to mind those of pinnules of adult individuals. They consist of twenty-five podia arranged in five radial triplets alternating with five interradial pairs. They are similar to the pinnular feeding structures of adults in that they have both collecting (radial) and paddling (interradial) podia.     

Fine-scale observations of the predatory behaviour of the carnivorous copepod <Emphasis Type="Italic">Paraeuchaeta norvegica</Emphasis> and the escape responses of their ichthyoplankton prey,Atlantic cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>)

Paraeuchaeta norvegica (8.5 mm total length) and yolk-sac stage Atlantic cod larvae (4 mm total length) (Gadus morhua) larvae were observed in aquaria (3 l of water) using silhouette video photography. This allowed direct observations (and quantitative measurement) of predator–prey interactions between these two species in 3-dimensions. Tail beats, used by cod larvae to propel themselves through the viscous fluid environment, also generate signals detectable by mechanoreceptive copepod predators. When the prey is close enough for detection and successful capture (approximately half a body-length), the copepod launches an extremely rapid high Reynolds number attack, grabbing the larva around its midsection. While capture itself takes place in milliseconds, minutes are required to subdue and completely ingest a cod larva. The behavioural observations are used to estimate the hydrodynamic signal strength of the cod larva’s tail beats and the copepod’s perceptive field for larval fish prey. Cod larvae are more sensitive to fluid velocity than P. norvegica and also appear capable of distinguishing between the signal generated by a swimming and an attacking copepod. However, the copepod can lunge at much faster velocities than a yolk-sac cod larva can escape, leading to the larva’s capture. These observations can serve as input to the predator–prey component of ecosystem models intended to assess the impact of P. norvegica on cod larvae.     

Apparent predation on ichthyoplankton by zooplankton and fishes in nearshore waters of southern California

Plankton collected at discrete depths in Santa Monica Bay, California, USA, during January 1982 were examined for fish eggs and larvae that had been attacked or consumed by zooplankton. The bongo net remained open for only 3 min and samples were preserved within 5 min of capture. Juvenile and adult fishes that had been captured by otter trawl and preserved within 20 min of capture were examined for ingested fish eggs and larvae. Three copepods (Corycaeus anglicus, Labidocera trispinosa, and Tortanus discaudatus), one euphausid larva (Nyctiphanes simplex), one amphipod (Monoculoides sp.), and an unidentified decapod larva were found attached to fish larvae in the preserved plankton samples (attachment to 23% of the fish larvae was observed in one sample). Overall, about 5% of the white croaker (Genyonemus lineatus) larvae and 2% of the northern anchovy (Engraulis mordax) larvae had attached zooplankton predators. Most fish larvae with attached zooplankton predators were small. We found no indication of zooplankton predation on fish eggs. Few fish eggs and larvae were found in the digestive tracts of juvenile or adult fishes, and the ingested fish larvae were relatively large. The discussion considers apparent preyspecificity of the zooplankton predators as well as potential biases that may be associated with preserved samples collected by nets.     

The effects of zooplankton swimming behavior on prey-capture kinematics of red drum larvae, <Emphasis Type="Italic">Sciaenops ocellatus</Emphasis>

Most marine fishes undergo a pelagic larval phase, the early life history stage that is often associated with a high rate of mortality due to starvation and predation. We present the first study that examines the effects of prey swimming behavior on prey-capture kinematics in marine fish larvae. Using a digital high-speed video camera, we recorded the swimming velocity of zooplankton prey (Artemia franciscana, Brachionus rotundiformis, a ciliate species, and two species of copepods) and the feeding behavior of red drum (Sciaenops ocellatus) larvae. From the video recordings we measured: (1) zooplankton swimming velocity in the absence of a red drum larva; (2) zooplankton swimming velocity in the presence of a red drum larva; and (3) the excursion and timing of key kinematic events during prey capture in red drum larvae. Two-way ANOVA revealed that: (1) swimming velocity varied among zooplankton prey; and (2) all zooplankton prey, except rotifers and ciliates, increased their swimming velocity in the presence of a red drum larva. The kinematics of prey capture differed between two developmental stages in S. ocellatus larvae. Hyoid-stage larvae (3–14 days old) fed on slow swimming B. rotundiformis (rotifers) while hyoid-opercular stage larvae (15 days and older) ate fast moving A. franciscana. Hyoid-opercular stage red drum larvae had a larger gape, hyoid depression and lower jaw angle, and a longer gape cycle duration relative to their hyoid-stage conspecifics. Interestingly, the feeding repertoire within either stage of red drum development was not affected by prey type. Knowledge of the direct relationship between fish larvae and their prey aids in our understanding of optimal foraging strategies and of the sources of mortality in marine fish larvae.     

Particle interception,transport and rejection by the feather star Oligometra serripinna (Echinodermata: Crinoidea), studied by frame analysis of videotapes

Suspension feeding by a stalkless crinoid (Oligometra serripinna) was studied at Lizard Island, Australia, in 1985. The crinoids were placed in a laboratory flume with a slow, unidirectional current of seawater. Nutritive and non-nutritive particles (15 to 180 m) were introduced upstream from the crinoid, and feeding behavior was recorded at high magnifications on videotape for frame analysis. These direct observations showed that each intercepted particle (whether a dejellied clam egg, Sephadex bead or latex sphere) contacts a single, evidently adhesive tube foot and is rapidly transferred to the pinnular food groove by a bend of the tube foot. The tube foot bends in about 0.1 s and returns to its extended position in 1 to 2 s. Spheres less than 20 m in diameter cause only the intercepting tube foot to bend. In contrast, larger spheres cause the coordinated bending of the intercepting tube foot plus many of the neighboring tube feet: the stimulus spreads through the reacting group of tube feet at about 1 cm s^-1. After transfer to the pinnular food groove, the nutritive particles (dejellied clam eggs) travel at about 1 cm min^-1 to the arm axis and thence down the arm food groove at about 4 cm min^-1 to the mouth; in contrast, non-nutritive particles (Sephadex beads and latex spheres) are discarded from the pinnular food groove between 1 and 30 s after capture. Tube-foot bending is presumably triggered when arriving particles (whether nutritive or non-nutritive) are detected by sensory cells in the tubefoot epithelium: mechanoreception by itself appears sufficient to initiate bending, although chemoreception may modify the reaction. Then, soon after captured particles have been transferred to the pinnular food groove, the crinoid discards those judged unsuitable (probably by contact chemoreceptors in the food-groove epithelium). Clam eggs with intact jelly layers temporarily hang up on tube feet they contact and then float away in the curent: the jelly evidently interferes with mechanoreception and/or chemoreception by the tube-foot epithelium. Some previous studies of crinoid feeding have suggested that particles are trapped in extensive nets or strands of mucus: we found no evidence for this in O. serripinna, which captures particles predominantly be the direct interception method of the aerosol filtration model.     

Alternative feeding mechanisms in megalopae of the blue crab Callinectes sapidus

The apparent mismatch between the energy requirements for planktotrophic growth and prey availability has long been paradoxical. One hypothesis to explain this paradox is that planktotrophic larvae display plasticity in feeding mechanisms in response to variable prey types and concentrations. This hypothesis was tested by videotaping megalopae of the brachyuran crab Callinectes sapidus Rathbun feeding on various-sized prey. Frame-by-frame analysis of the videotapes indicated that C. sapidus megalopae used both raptorial and suspension feeding to capture prey while in the water column. Raptorial feeding was used to capture macro-zooplankton, including copepods. The swimming form of suspension feeding was based on a modified fling-and-clap mechanism using the chelipeds. Suspension feeding while at rest utilized a weak current generated by the mouthparts to direct prey to the mouth. Both suspension-feeding mechanisms resulted in the efficient capture of rotifer-sized particles. The energy/handling time ratios for all three feeding mechanisms are very similar (E/H range 0.016-0.019 µg C s^-1) for the natural prey tested. These results support the hypothesis that feeding in brachyuran larvae is plastic and includes mechanisms of both raptorial and suspension feeding. The ability to suspension feed at rest is adaptive, since megalopae use selective tidal transport to re-invade an estuary and may spend up to 18 h day^-1 clinging to a benthic substrate. Electronic supplementary material to this paper can be obtained by using the Springer LINK server located at http://dx.doi.org/10.1007/s00227-002-0781-1.     

Influence of flow speed on the functional response of a passive suspension feeder,the spionid polychaete <Emphasis Type="Italic">Polydora cornuta</Emphasis>

Passive suspension feeders rely on surrounding flow to deliver food particles to them. Therefore, the classic conception of functional response (feeding rate vs. food concentration) may require modification to account for flow speed as a second independent variable. I compared the functional response of Polydora cornuta at different velocities and determined whether food capture was proportional to particle flux (concentration × velocity). To understand feeding responses at a mechanistic level, I measured the functional responses in terms of contact and capture rates and determined particle retention efficiency. Experiments were run separately with two sizes of food particles, and with juvenile or adult worms. For both worm sizes and both particle sizes, capture rate in weak flow was directly related to concentration, but in strong flow it was constant. Worms were therefore unable to benefit from abundant food when in strong flow. The critical velocity at which the capture rate became constant was lower for adult worms than for juvenile worms, and it was lower for small particles than for large particles. Retention efficiency was constant among all treatments, and the results for contact rate were essentially the same as for capture rate. Therefore, the mechanics of particle contact must explain the effects of velocity on the functional response. Contact rate was not a constant proportion of particle flux; treatments with similar fluxes yielded different contact rates depending on the strength of flow. The results appeared to be caused by a velocity-induced behavioral change in appendage posture that affects contact rates: in moderate flow, worms form their feeding palps into helical coils, which they tighten as the velocity increases. I suggest this behavior constrains suspension feeding rates and the mechanical selection between particle sizes when worms are in strong flow, and that the effect changes with ontogeny. Because the results are consistent with patterns in measured growth rates of P. cornuta, I hypothesize that this influence of velocity on the functional response can constrain growth and population dynamics in this species.     

Analysis of capture efficiency in suspension feeding: Application of nonparametric binary regression

Suspension feeding is an important mechanism by which many marine organisms obtain food. A nonparametric method for analyzing experimental results on capture efficiency in suspension feeding is described. This approach can be used to estimate capture efficiency along the length of a cilium and to compare the capture efficiencies of two different cilia. The method is applied to experimental data on feeding by larvae of the clamMerenaria merenaria (Linné), to examine changes in capture efficiency through time, and to assess the effect of particle electrostatic charge on capture efficiency.     

Functional morphology and feeding behavior of Scolelepis squamata (Polychaeta: Spionidae)

Functional morphology and feeding behavior of Scolelepis squamata (Müller) were studied. Gut contents consisted of unconsolidated sedimentary particles, fecal pellets of other species, and a wide variety of embryos, larvae, and juveniles. Unlike other spionid polychaetes the palps of S. squamata lack a median, ciliated groove. Particles captured by the palps were brought to the pharynx by a complete contraction of the palp. In the presence of a current, S. squamata helically coiled their palps, and fed almost exclusively on suspended and resuspended particles. In contrast to most spionid polychaetes the palps of S. squamata are not deciduous. Various aspects of the morphology and feeding behavior of S. squamata are discussed in terms of their functional and ecological significance.     

Dietary study of the midwater polychaete Poeobius meseres in Monterey Bay,California

This study presents the first quantification of the diet of a gelatinous midwater organism on a temporal basis. Using the Monterey Bay Aquarium Research Institute's remotely operated vehicle Ventana, regular collections of the polychaete Poeobius meseres (Heath, 1930) over a 1 yr period (October 1990 to November 1991) in Monterey Bay yielded intact organisms for the study of feeding behavior and quantitative analysis of stomach contents. In situ observations showed P. meseres feeding in two different ways: (1) by deployment of a mucus web in the water column that passively collects particles for consumption; and/or (2) by grasping detrital material in the water column with its ciliated tentacles. Stomach-content analyses showed that P. meseres is primarily coprophagic, its diet being dominated by fecal pellets from euphausiids and copepods. These fecal pellets appear to provide P. meseres with essentially all its carbon. Although fecal pellets were the most important food item volumetrically, P. meseres also consumed large numbers of diatoms and small numbers of dinoflagellates, chrysophytes, radiolarians, foraminiferans and eggs. The diet of P. meseres appears to reflect primary productivity in the surface waters, with different food items predominant in the diet at different times of the year. Pennate diatoms were most abundant in the diet during the fall, centric diatoms were most abundant during the sumnier, and fecal pellets during the winter. The composition of P. meseres diet suggests that this and other midwater gelatinous organisms have a significant role in the remineralization of particles as they sink from the surface to the deep sea.     

Suspension feeding in the brittle-star Ophiothrix fragilis : efficiency of particle retention and implications for the use of encounter-rate models

Dense beds of the suspension-feeding brittle-star Ophiothrix fragilis are common in European waters. Their potential importance in benthic–pelagic coupling has been highlighted, but little is known about the feeding dynamics of this species. Encounter-rate models provide a potential mechanism for the estimation of feeding rates on suspended material of varying sizes. This work investigates factors essential to the application of such models. Particle-retention efficiency (RE) converts encounter rate into capture, or clearance rate. Laboratory studies demonstrated that RE varied with the interactive effects of flow velocity and particle size. RE was lowest for large particles, particularly at high flow velocity where RE as low as 59% was observed. This indicates that if RE is not accounted for in encounter-rate models, significant overestimates of feeding rates on large particles may occur. Flow around feeding arms and tube feet was characterised by intermediate Reynolds numbers, precluding application of the most simple encounter-rate models. Complex secondary-flow patterns were observed, which carried particles along the downstream side of the feeding arms, but these did not appear to increase the area available for particle capture. Previously reported particle capture by arm spines was not observed. Evidence of active rejection of large particles by tube feet was recorded. Difficulties in the application of encounter-rate models for prediction of seston-removal rates are highlighted by these results. Predicted encounter rate may deviate from actual clearance rate due to the effects of retention dynamics, localised flow patterns and differential particle handling. Other methods of estimation of seston-removal rates are equally problematic however, so that encounter-rate models are likely to remain a useful tool for such estimates. Received: 23 January 1998 / Accepted: 24 June 1998     

On the feeding and behaviour of the larva of Branchiostoma lanceolatum

Collections of Branchiostoma lanceolatum (Pallas) made in mid-May and mid-July at Helgoland before and after spawning have established that the larvae leave the amphioxus ground about June and therefore presumably become planktonic. Metamorphosing larvae and young adults can be collected on the ground in late August and early September and are either the same larvae returning, or others from a neighbouring ground within the same circulating current system. An examination of the gut contents of 67 larvae collected from the plankton at Helgoland in August showed that 30% of the animals had ingested calanoid copepods or other organic material of a size similar to that of the larval mouth. A few larvae had also taken small particles evidently by a ciliary mechanism. In 50% of the larvae the gut was empty. It has been found that, in addition to a muscular mouth and gill bars richly supplied with nerves, both the gut wall and the body wall are muscular and capable of passing, by peristalsis, large food masses that distend the body. The visceral muscles of the larva resemble the coelomyarian fibres of the Nematoda. The larva appears, therefore, to be both microphagous and macrophagous. Evidence from the swimming behaviour and from reports of the vertical distribution of larvae in the sea is discussed. It is suggested that the larvae normally swim upward with the mouth and gills closed and then sink passively in the horizontal position with the pharynx expanded and the open mouth directed downward. In the event of large organism such as a copepod or a mass of organic material coming into contact with the adhesive lower left surface of the larva, it could be captured by the mobile lower lip and engulfed. The straightening of the larval tail, the great increase in the number of eyecups and the growth of the metapleura at metamorphosis are suggested as factors leading to the settlement of the young adult. Attention is drawn to the possible significance of the structure of the larva in interpreting the relationships of the cephalochordates.     

The role of mucus in particle processing by suspension-feeding marine bivalves: unifying principles

Contemporary research on bivalve suspension-feeding has revealed a diversity of particle processing mechanisms depending on the anatomy and functioning of the pallial organs involved. On the biochemical level, however, some evidence of homogeneity has emerged concerning the types of mucopolysaccharide associated with particle processing. The present study uses both previous data and original research combining video endoscopy and mucocyte mapping to further explore the relationships between pallial organ topography, functional correlates, direction of current flow, and mucocyte secretion type. Five species representing five different families and all four major gill types are represented: Mytilus edulis, Placopecten magellanicus, Crassostrea virginica, Mya arenaria, and Spisula solidissima. Viscous acid or acid-dominant mucopolysaccharides are used when particle transport occurs on an exposed surface, or on a structure leading directly to such a surface, counter to the prevailing current flow. Associated functions are indiscriminate transport in gill ventral particle grooves and rejection of pseudofeces. Lower-viscosity mixed mucopolysaccharides are used when particle transport is on an enclosed or semi-enclosed surface, leading to other such surfaces, and with the current flow. Associated functions are transport of particles destined for ingestion, and ingestion itself. Low-viscosity neutral mucopolysaccharides are found in regions where reduction of mucus viscosity is important, such as the areas of the labial palps responsible for fluidization of the high-viscosity mucus-particle cord of the gill ventral particle groove prior to particle extraction. There thus appears to be a specialization of mucus type corresponding to functional specialization of the various pallial organs in suspension-feeding marine bivalves. Received: 10 October 1996 / Accepted: 7 May 1997     

New observations of the gills of Placopecten magellanicus (Mollusca: Bivalvia), and implications for nutrition

The organization, general anatomy, and surface microanatomy of all regions of the gills of a representive bivalve mollusc, Placopecten magellanicus Gmelin, were studied using stereo-microscopic, histological, and scanning electron microscopic techniques. Individuals were collected in May and November 1985 from Chamcook Bay, New Brunswick, Canada. In addition to correcting earlier accounts of this structure, a number of new observations are reported. The orientation of the ciliated spurs appears to be responsible for the sinusoidal arrangement of the gill filaments. Micrographs showing the structure of the dorsal respiratory expansion are presented. The entire abfrontal surface of the principal filament, including the dorsal respiratory expansion, is densely ciliated and mucosecretory. These characteristics may aid in the establishment of a respiratory current and in the prevention of gill damage during escape responses. All nonciliated regions of the gill filaments are covered with microvilli, thus greatly increasing the surface area of the gill. The feeding mechanism is discussed in relation to the dorsal and ventral ciliated tracts. Symbiotic ciliate protozoans are constantly dislodged from the gill filaments and transported via the ventral mucus string to the buccal region. The nutritional implications of these observations are discussed.     

Embryonic,larval, and post-metamorphic development of the sea urchin <Emphasis Type="Italic">Cassidulus mitis</Emphasis> (Echinoidea; Cassiduloida): an endemic brooding species from Rio de Janeiro,Brazil

Cassiduloids are currently rare irregular echinoids with a highly conserved adult morphology. Aristotle’s lantern is present only during the post-metamorphic stage, and little is known about the early development of species in this group. Cassidulus mitis produces eggs of about 375 μm in diameter, lecithotrophic larvae with four reduced arms with skeletal fenestrated rods, cilia along the body surface, and a ciliated band on arms and lobes. Offspring is brooded among the female spines from embryo to settler’s stage. The echinopluteus larval stage is reached 6 days after fertilization, and the settler’s stage is formed at the age of 17 days. Aristotle’s lantern appears around the thirteenth day of development. The lantern is well developed and functional in settlers. It remains until at least 62 days after fertilization and can be used to acquire food from the environment. The early development of C. mitis is unusual concerning features of typical lecithotrophic larvae (such as reduced arms), but retains some features of planktotrophic larvae (such as skeletal rods and a ciliated band). Regarding egg size, early development in C. mitis seems to be transitioning from facultative lecithotrophic to typical obligate lecithotrophic pattern in echinoid larval evolution.