Prey Capture in Ly Onsiella Form Osa (Bivalvia: Anomalodesmata
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Pacific Science (1984), vol. 38, no. 4 © 1985 by the University of Hawaii Press. All rights reserved Prey Capture in Lyonsiella formosa (Bivalvia: Anomalodesmata: Verticordiacea) 1 BRIAN MORTON2 ABSTRACT: A specimen of the bathyal verticordiid Lyonsiella formosa has been obtained from Hawaii at 460 m depth. Assignment of this specimen to L. formosa suggests that this species has a much wider range than hitherto believed. Dissection and subsequent histological examination ofthe specimen suggests a mechanism of prey capture completely different from that previously described for this species and resembling that attributed to Poromya granulata. Sensory papillae on the siphonal tentacles probably detect the prey. Prey capture is by eversion of an enormous hoodlike cowl of the inhalant siphon. Inversion brings the prey into the mantle cavity. Further distension of the siphon within the mantle cavity is believed to push the prey into the buccal apparatus comprising medially fused labial palps . The unfused tips of the palps or the foot may assist in this. A model of the hydraulic changes that may occur in Lyonsiellaformosa to effect prey capture is described. The similar modes offeeding exhibited by L.formosa (Verticordiidae) and Poromya granulata (Poromyidae) suggest a close affinity. ON TH E DEEP OCEANFLOOR two bivalve taxa the mechanism of prey capture. Feeding in have adopted a predatory mode of life. Pos- . Poromya was explained by Morton (1981a) sibly all species of the Propeamussiidae are who showed that the inhalant siphon com carnivores (Knudsen 1970, 1979), though the prised a huge raptorial cowl beneath which mechanism of prey capture is unknown. prey were trapped before being withdrawn Con versely, the Parilimyidae, Verticordiidae, into the mantle cavity. Morton (1981b) also Poromyidae, and Cuspidariidae of the ancient explained prey capture in Parilimya , the rap and peculiar subclass Anomalodesmata have torial inhalant siphon this time being with well-known predatory habits. Each species drawn by elongate siphonal (taenioid) retrac has particular morphological adaptations to tor muscles. Allen and Turner (1974) investi its predatory life style, but the single most im gated the Verticordiidae in great detail but did portant feature of them all is the active mus not come to any firm conclusions with regard cular pumping of fluids in and out of the to how prey is captured, stating that (p. 513) mantle cavity, to facilitate prey capture, in "the tentacles in L. abyssicola and probably in stead of the ciliary movement of fine sus all other species, extend across the surface pended material. The prey is digested by pro layer of the abyssal sediment in life and pas teolytic enzymes (Reid 1977) within a highly sively wait for organisms to brush against them modified gut (Purchon 1956). Yonge (1928) and adhere to them"; and later (also p. 513) investigated Poromya and Cuspidaria but " the tentacles with adhering food contract Reid and Reid (1974) were the first to demon and move inwards into the inhalant aperture strate the mechanism of prey capture in Cuspi where they are wiped clean by the constriction daria. This was subsequently confirmed by of the inhalant aperture and valve-when the Allen and Morgan (1981) who also investi latter is present. " This view was later re gated poromyids but could not elaborate on iterated by Allen (1983). Allen and Turner (1974) describe a variety of siphonal types for the verticordiids they in 1 Manu script accepted II April 1984. 2 The University of Hong Kong, Depart ment of Zoo vestigated, suggesting a greater diversity of logy, Hong Kong. feeding strategies than sticky tentacles. Ac- 283 284 PACIFIC SCIENCE, Volume 38, October 1984 cordingly, Lyonsiella formosa is here investi a box dredge operating at a depth of 460 m gated in greater detail, special attention being and with a sea temperature of 9° C. No ob paid to the possible mechanism of prey servations on the living animal were possible, capture. as it was dead by the time the dredge reached the surface. It was preserved in 5 percent neu tral formalin. Following dissection, the speci men was transversely sectioned at 6 Jlm and TAXONOMY alternate sections stained in either Ehrlichs' Lyonsiellaformosa (Jeffreys, 1881)has been haematoxylin and eosin or Masson's tri further described by Allen and Turner (1974) chrome. The broken shell has been deposited who also illustrated the structure of the body in the collections of the Bernice P. Bishop as well as the shell. The species has only been Museum (Reg. no. 207491). recorded from the Atlantic (i.e., the Canaries, Azores, Bay of Biscay, and GulfofMexico) at depths ranging between 366 and 3783m (Allen and Turner 1974, Knudsen 1979). I have also FUNCTIONAL MORPHOLOGY examined the type specimen (AMNH Reg. no. The Shell 61238). Lyonsiella elegans (Thiele and Jaeckel, 1931) The shell (Figure lA) is exceedingly fragile, has only been recorded from the type locality , equivalve, and markedly inequilateral. The Station 242 (404 m) of the Valdivia Expedition anterior face is rounded and somewhat re (1898- 1899) (6° 34.8' S, 39° 35.5' E) at Dar es duced relative to the posterior which is in Salaam. I have not seen the type specimen of flated. The posterior margin is ventrally this species, there being an excellent illus angular and has a corrugated outline. tration of it (Thiele and Jaeckel 1931). Whereas the anterior face is smooth, the pos Thiele and Jaeckel acknowledge that Lyon terior is radially ridged and concentrically siella elegans resembles the Atlantic L. for ringed. From each umbone arises a single rib mosa, and Allen and Turner (1974) have that extends to the ventral margin just pos shown that the shells of different -sized speci terior to the mid line. Seven or eight further mens ofthe latter species vary considerably in rays extend to the posterior and posterodorsal overall form and in details of the sculpture. margin of the shell. The specimen here under consideration was At the junction of each ray with a con obtained from a depth of 460 m off Hawaii centric ring a sharp spine is produced, though and bears a very strong resemblance to L. in this specimen most have been broken off. elegans. Bearing in mind the great variability Spines in different positions adorn the shells in shell form of L.formosa, however, and as ofLyonsiellaformosa illustrated by Allen and suming that L. elegans is possibly equally vari Turner (1974), but a series of spines on the able, it seems very probable that L. elegans is dorsal part of the shell appear more resistant but ajunior synonym of L.formosa. As will be and characterize all individuals, including the seen later, the specimen is morphologically type of L. elegans (Thiele and Jaeckel , 1931). indistinguishable from L. formosa. It accord The shell between the posterior rays and the ingly seems likely that L. formosa is not re single more median rib is concave. Allen and stricted to the Atlantic, but is a widely dis Turner (1974) illustrate exceedingly fine rays tributed bathyal species. on the anterior face, these are barely discern ible in this specimen . The surface of the shell is finely grained. From the dorsal aspect (Fig ure 1B) the great expansion of the posterior MATERIALS AND METHODS region of the shell is more clearly seen. This re The specimen of Lyonsiellaformosa here in sults from , as will be seen later, the location of vestigated was obtained during a short re the prey-capturing organs posteriorly. The search visit to Hawaii in December 1981 with shell is widely emarginate posteriorly and less Prey Capture in Lyonsiella-MoRToN 285 L 2·5mm c lmm F IGURE I. LyonsiellaJormosa. The shell as seen from A, the right side, and B, the dorsal aspect; C, an enlarged view of the hinge plate of the left valve. FP, fused pcriostracum; ILL, inner ligament layer; L, lithodesma. so anteriorly. The umbones point slightly tiny pedal retractor muscles-anterior (APR) forward. and posterior (PPR). Other muscles are con Hinge structure has similarly been described cerned with pallial retraction (Figure 7A, by Allen and Turner (1974) (Figure IC). PRM). From the exhalant siphon extends a There are no hinge teeth. The primary liga thin array of muscles (Figure 2, ESR) which ment is small and internal, comprising inner posteriorly attach to the shell approximately ligament layer (ILL) only, this being calcified where the dorsal margin ofthe ascending lam mid-ventrally into an inconspicuous litho ella of the outer demibranch unites with the desma (L). The dorsal borders of the shell are mantle. united by a thin layer of fused periostracum The inhalant siphon is retracted by a pair of (FP) forming a " secondary" ligament whose muscles within the fused ventral mantle mar main function is to assist in the alignment of gin and which have their origins posterior to the valves. the pedal gape. The muscles are attached at the single near-median rib on each valve and extend posteriorly into the inhalant siphon, The Musculature radiating as they do. The two adductor muscles (Figure 2, AA; Since the muscles are probably derived PA) are of approximately equal size (iso from the more typical bivalve siphonal retrac myarian) and are bordered internally by very tor system, these are called taenioid muscles 286 PACIFIC SCIENCE, Volume 38, October 1984 PPR PA 00 ES ESR APR H-- - AA FLP J....'...'.'..'. Zmrn TM < FO PPG BG T BYG FIGURE 2. Ly onsiellafo rmosa. The organs of the mantle cavity after removal of the right shell valve and mantle lobe. AA , anterior adductor muscle; APR, anterior pedal retractor muscle; BG, byssal gland ; BYG, byssal groove; DD, digestive diverticula; ES, exhalan t siphon; ES R, exhalant siphonal retractor muscle; FLP, fused labial palps; FO, foot; HE, heart; ID, inner demibranch; K, kidney; LS, lacunal system; 0 , ovary; OD, out er demibranch; P, periostracum; PA, posterior adductor muscle; PE, pericard ium; PPG, posterior limit of pedal gape; PPR, posterior pedal retractor muscle; SC , siphonal cowl; T, testis; TM, taenioid muscle.