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• W2330556182 abstract "Particle capture, transport, ingestion, and selection mechanisms used by Dipolydora commensalis, a symbiotic worm that inhabits burrows it produces in hermit crab shells, were studied in the laboratory. Worms removed from shells inhabited by Pagurus longicarpus were placed in capillary tubes for observation and video-microscopy of feeding. Particles were captured by the palps and transported to the mouth by ciliary and muscular action. Mean transport rates for cysts of Artemia from the distal ends of the palps to the mouth ranged from 0.7 to 1.3 mm/sec; those for glass beads, from 0.2 to 0.8 mm/sec. Upon reaching the mouth, particles were engulfed by the lateral lips and ingested. Worms feeding on large particles (>0.2 mm) repeatedly arched the anterior end of the body. Nauplii of Artemia were used to determine the ability of the worms to feed on small motile organisms over a 24-h period in the dark. In six trials the percentage of nauplii ingested ranged from 58% to 76%. In particle selection experiments, worms were offered uncoated glass beads and beads coated with fluid from the ribbed mussel, Geukensia demissa. In two experiments 91% and 89% of coated glass beads were ingested as compared to 15% and 4% of uncoated beads, respectively. Video-analyses of feeding sequences showed that cysts were often partially ingested and then ejected from the mouth. Uncoated beads transported along the palps often never reached the mouth. These results indicate that D. commensalis can actively select particles at the mouth and possibly during transport along the palps. Additional key words: suspension feeding, deposit feeding, feeding biodynamics, videoanalysis, Annelida Polychaetous annelids of the family Spionidae occupy a variety of marine and estuarine habitats, ranging from soft-bottom sediments to hard calcareous substrates (Fauchald & Jumars 1979). The biology of shell-boring spionids has been studied in considerable detail, in part due to their effects on commercially important species of bivalves (e.g., Haigler 1969; Blake & Evans 1973; Zottoli & Carriker 1974; Sato-Okoshi & Okoshi 1993). Blake (1996) referred those species in the genus Polydora bearing notosetae on setiger 1 to the genus Dipolydora VERRILL 1879, making the current combination for the species under study Dipolydora commensalis (ANDREWS 1891). Although many spionids are non-specific borers, D. commensalis burrows exclusively into gastropod shells inhabited by hermit crabs. The burrows of the worm often begin on the columellar side of the shell aperture and extend to a Author for correspondence. E-mail: jwil4024@uriacc.uri.edu the internal apex of the shell via a thin secreted calcareous tube (Andrews 1891; Hatfield 1965; Radashevsky 1989). D. commensalis is usually classified as a commensal, but recent research suggests that the nature of this association with hermit crabs may require some re-evaluation. Buckley & Ebersole (1994) showed that the strength of shells inhabited by spionids is reduced, decreasing the resistance of shells to crushing forces applied by predators. The feeding biology of polychaetes has been summarized by Fauchald & Jumars (1979) and several spionids have been investigated in detail (Dorsett 1961; Dauer et al. 1981; Levin 1981; Jumars et al. 1982; Dauer 1983, 1984, 1985; Yokoyama 1988; Miller et al. 1992; Bock & Miller 1996; Shimeta & Koehl 1997). In most spionids, a pair of peristomial palps are used to capture and transport food particles for ingestion. Particles are usually transported, by a combination of muscular movement and ciliary action, in a median ciliated groove (Dorsett 1961), although in at This content downloaded from 157.55.39.186 on Sun, 09 Oct 2016 06:17:45 UTC All use subject to http://about.jstor.org/terms Williams & McDermott least two species the palps lack a ciliated groove (Dauer & Ewing 1991). Spionid polychaetes have been classified as either suspension-feeders or deposit-feeders, or as suspension-deposit feeders (Dauer et al. 1981). Suspensiondeposit feeding spionids have a mixed mode of feeding, depending on environmental conditions, and change their feeding behavior in the presence of particle fluxes or deposited material (Taghon et al. 1980; Miller et al. 1992; Bock & Miller 1996). Particle capture in spionids occurs primarily through direct interception of food by the palps, although inertial impaction and gravitational deposition may have slight effects (Taghon et al. 1980; Shimeta & Koehl 1997). Several physical limits may affect the ability of a worm to feed on particles. Mucus secreted at the distal end of the palps and along the ciliary oral groove has a limited strength of adhesion to intercepted particles (Jumars et al. 1982). There are also limits to the size of particles that can be transported along the palps, via ciliary and/or muscular action. The size of the mouth and/or pharynx acts as the final determinant of the size of particles that can be ingested. Most studies on the feeding biology of spionids have involved species inhabiting soft-bottom substrates. The use of soft-bottom spionids has allowed investigators to study the feeding biology in a variety of contexts (e.g., Dauer et al. 1981; Jumars et al. 1982; Taghon 1982; Taghon & Jumars 1984; Shimeta 1996), in particular, active and passive particle selection. Active particle selection in spionids occurs at the mouth, pharynx, or along the ciliated oral groove of the palps (Dauer et al. 1981; Levin 1981; Shimeta & Koehl 1997). Passive selection results from particle loss due to limited adhesive strength of mucus on the palps (Self & Jumars 1978; Taghon 1982; Shimeta & Koehl 1997). Such particle loss can occur at the point of initial contact between palp and particle or during particle transport along the palp (Taghon 1982). Size, specific gravity, and texture of food particles can affect the passive selection mechanisms in spionids (Self & Jumars 1978; Jumars et al. 1982; Taghon 1982; Shimeta & Koehl 1997). Passive selection mechanisms, therefore, can cause preferential ingestion of particles that are smaller, rougher, and have lower specific gravities. Passive loss could also determine the ability of a polychaete to engage in a macrophagous, predatory mode of feeding documented in certain spionids (Dorsett 1961; Jumars et al. 1982). One might expect differences between the feeding behavior of soft-bottom species and D. commensalis due to the distinct habitat of this species. The burrow in which D. commensalis lives may prevent it from feeding at the sediment-water interface. Additionally, interactions with the hermit crab host may influence the feeding behavior of the worm, which has been reported to feed on fine particles suspended in the branchial currents of the crab (Dauer 1991; Radashevsky 1993), on particles attached to the legs of the crab (Dauer 1991), and on nauplii of Artemia being fed to the longwrist hermit crab Pagurus longicarpus SAY 1817 (J.J. McDermott, unpubl.). Since the original description of D. commensalis, only nonquantitative obs vations on the feeding behavior of this polychaete have been reported (Radashevsky 1993); however, palp morphology and its relation to feeding mechanisms on small particles was studied by Dauer (1991). The objective of our study was to investigate the feeding biology of D. commensalis, focusing on mechanisms of particle capture, transport, ingestion, and selection, by combining observations of behavior with quantitative measurements of deposit and suspension feeding. The observation of feeding behavior and measurements of feeding rates were made possible by removing the worms from the shells. Worms placed in glass tubes were compared with those in shells inhabited by P. longicarpus, as they manipulated cysts and swimming nauplii of Artemia and glass beads. Videomicroscopy, a valuable aid in studying the feeding behavior of small invertebrates and their larvae (e.g., Strathmann 1982; Gilmour 1985; Holland et al. 1986; Gallager 1988; Emlet 1990; Hansen & Ockelmann 1991; Nielsen et al. 1993; Mayer 1994; Chen et al. 1996; Hart 1996; Shimeta & Koehl 1997), was used to record the feeding rates of the worms and mechanisms involved in feeding." @default.
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• W2330556182 title "Feeding Behavior of Dipolydora commensalis (Polychaeta: Spionidae): Particle Capture, Transport, and Selection" @default.
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