<I>Sabellastarte Magnifica</I>
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FEEDING AND TUBE-BUILDING IN SABELLASTARTE MAGNIFICll (SHAW) (SABELLIDAE:POLYCHAETA) GAIL FITZSIMONS Bellairs Research Institute of McGill University, St. James, Barbados, West Indies ABSTRACT The branchial crown of Sabellastarte magnifica (Shaw) acts as a filtra- tion device for trapping particles in suspension in the sea water. The anatomy of the branchial crown, the mechanism of collecting and sorting particles, and the method of utilizing the particles for feeding and tube- building are studied. The general chemical nature of the tube is described. INTRODUCTION Sabellastarte magnifica (Shaw) is a common West Indian littoral Sabel- lid polychaete. The literature on this species is restricted almost exclusively to faunal records and descriptions of the external morphology. Sabellastarte magnifica has been recorded from the Bahamas (Andrew & Andrew, 1953), Jamaica (Shaw, 1800; Schmarda, 1861; Augener, 1922; Marsden, 1960), Haiti (Augener, 1922), St. Thomas (Augener, 1922, 1925), St. Croix (Augener, 1922, 1925), Antigua (Mullin, 1923; Treadwell, 1924), Guadeloupe (Kinberg, 1867), Barbados (Augener, 1922; Marsden, 1960), Isla de Margarita (Monro, 1933), and Cura<;ao (Augener, 1927). It has also been found along eastern Mexico (Augener, 1922; Rioja, 1946; Hartman, 1951), on the Caribbean coast of Panama (Monro, 1933), and on the northern coast of South America, off Venezuela and Surinam (Augener, 1922). Most of the faunal records noted above include descriptions of the external morphology. Mullin's (1923) paper contains the only report on internal anatomy to be found in the literature on this species. She describes the digestive system, the septa, and the muscular, circulatory, nervous, and excretory systems. Mullin was not able to identify the reproductive system and concluded that it probably consists of gonads that are inconspicuous except during breeding. The only detailed account of feeding and tube-building in any Sabellid is found in Nicol's (1930) paper on Sabella pavonina. Comparisons will be drawn between Nicol's description of Sabella pavon ina and the obser- vations on Sabellastarte magnifica. The nomenclature of Nicol has been followed in considering the orientation of the filaments and pinnules. The surface toward the centre of the branchial funnel is designated as the frontal face, that towards the outside of the funnel the abfrontal face, and those to the sides the lateral faces. Sabellastarte magnifica belongs to the subfamily Sabellinae of the family Sabellidae and is morphologically a typical Sabellid. The typical full-sized 1965] Fitzsimons: Feeding and Tube-Building in Sabellastarte magnifica 643 Sabel/astarte magnifica in Barbados is about 130-160 mm long, including the branchial crown. The branchial crown usually constitutes one-half to three-quarters of the total body length. The crown spread is approximately 102-127 mm. The body is thickset, about 12 mm across and of uniform width, except at the posterior end where the last 20-25 segments taper to a blunt point. The body is dorsally convex and ventrally flattened. There are 8 thoracic segments. The number of abdominal segments is variable, usually about 125 to 150. As in all Sabellids, the two processes of feeding and tube-building are closely interrelated in Sabel/astarte magnifica. For both processes the worm utilizes particulate material in suspension in the surrounding sea water. The branchial crown is the particle-collecting mechanism. The work was supported by a grant from the National Research Council of Canada. I am grateful to Dr. John B. Lewis for his advice and encour- agement. EXPLANA nON OF LETTERING A.-abfrontal cilia folds A.C.-anastomosing cells in skeletal I.B.P.-inner edge of basal piece sheath J.-bending joint of pinnule B.D.L.-base of dorsal lip L.-Iatero-frontal cilia B.F.-basal fold L.BR.F.-Iower branchial filament B.G.-basal groove L.C.G.F.-Iongitudinal ciliated groove B.GR.-basal granules of filament B.L.L.-base of lateral lip L.L.-Iateral lip BR.F.-branchial filament L.M.P.-longitudinal muscles of B.V.BR.F.-blood vessel of branchial pinnule filament L.R.B.F.-longitudinal ridge of B.Y.P.-blood vessel of pinnule basal fold B.Y.PP.-blood vessel of palp L.R.D.L.-Iongitudinal ridge of B.W.-basal web dorsal lip C.-canal for basal branchial blood L.R.L.L.-Iongitudinal ridge of vessel lateral lip C.G.M.-ciliated groove leading to M.-mouth mouth M.c.-mucous cell c.G.V.S.-ciliated groove leading to N.C.-nucleus of ciliated cell ventral sac N.E.C.-nucleus of epidermal cell CU.-cuticle N.L.M.F.-nucleus of longitudinal D.L.-dorsal lip muscle fiber D.P.-dorsal pit O.B.P.-outer edge of basal piece D.P.C.F.-dorsal peristomial collar P.-pinnule fold P.c.-penetrating cell of epidermis E.-epidermis P.F.-'-parallel fold E.C.-epidermal cell PP.-palp E.G.E.-elevated glandular epidermis R.D.L.-rim of dorsal lip F.-frontal cilia R.L.L.-rim of lateral lip F.G.P.-frontal groove of pinnule S.A.B.P.-skeletal axis of basal piece G.-groove between the bases of the S.A.BR.F.-skeletal axis of branchial branchial filaments and the lateral filament lip S.A.P.-skeletal axis of pinnule G.P.F.-groove between the parallel ,S.A.PP.-skeletal axis of palp 644 Bulletin of Marine Science [15(3) S.R.PP.-supporting skeletal rod ciliated groove of palp V.P.c.P.-ventral peristomial S.S.A.-sheath of skeletal axis collar fold U.BR.P.-upper branchial filament V.S.-ventral sac V.L.C.G.-ventrallongitudinal W.-web joining the paired palps FEEDING Methods.-lt was possible to keep Sabel/astarte magnifica, in good condition, for about two weeks in concrete laboratory aquaria through which sea water was continually circulated. After about two weeks, however, the worms began to lose their muscular tone and reactivity and to spontaneously autotomize their branchial crowns. Only specimens freshly taken from the sea were used in studying the feeding process. The most satisfactory method of study was found to be that used by Nicol for Sabella pavonina. The worm, still in its own tube, was placed inside a glass tube of slightly greater diameter. This was placed in a jar filled with fresh sea water and was supported and adjusted to any position desired by means of wire loops. The observations were made with a binocular dissecting microscope and also with a fluorescent lamp equipped with a magnifying glass window. The finer details of the ciliary tracts were studied by examining excised portions of the feeding apparatus placed in sea water under high power magnification. The ciliary tracts were also examined by staining unfixed portions of the feeding apparatus with Noland's (Pantin, 1948) stain which stains cilia bright pink. Noland's stain stopped the movement of the cilia. Small particles present in the sea water usually made the currents visible. When necessary, various particulate substances, such as carmine powder, Sira rouge powder, finely ground carborundum, sand grains, and starch grains stained with iodine, were placed on or near the branchial crown. In studying the anatomy of the branchial region, both fresh and pre- served specimens were used. The formalin-ethanol preservation method recommended by Marr (1962) for polychaetes was found to give the best results. The detailed structure of the crown was studied by cutting IOfL serial sections in the longitudinal and transverse planes. The material was fixed in Duboscq-Brasil (Nicol, 1930), embedded in paraffin and stained in Heidenhain's hematoxylin and eosin. Anatomy of the branchial region.- The branchial crown is very showy and occurs in a wide variety of colors. The branchial filaments are colored in shades of brown with several series of paler spots forming bands. The most frequent color combinations are fuscous brown banded with light tan spots, chocolate brown banded with white, dark purple with brown spots, and dark mahogany red banded with light brown. Some crowns are 1965] Fitzsimons: Feeding and Tube-Building in Sabel/astarte magnifica 645 A B.F. B.f o B c FIGURE 1. The branchial crown of Sabellastarte magnifica. A. Anterior view of the inside of the branchial crown with the filaments fully spread; the fila- ments have been cut off at their bases, the pinnules are not shown and the dorsal lips are shown pulled towards the mid-line. B. Ventral extremity of the right lateral lip. C. Frontal view of three branchial filaments; the pinnules of the lower filament are not shown. D. Abfrontal view of five branchial filaments. 646 Bulletin ot Marine Science [15(3) almost white with indistinct barring, like watered silk (Hartman, 1951). Figure lA shows an anterior view of the branchial region with the crown fully expanded. The prostomium is very compressed and bears the following structures on its anterior surface: two paired palps, two paired ventral sacs, a mouth, a pair of lateral lips, a pair of dorsal lips, and a branchial crown. The branchial crown consists of two semicircular stems, one on each side of the anterior face of the prostomium. The two stems are connected on the dorsal side but are separate on the ventral side. Anteriorly, the two stems branch out into numerous filaments. The filaments vary in number in different individuals but are approximately the same number on both sides of the crown. In a full-sized specimen there are about 50-60 filaments on each side. The filaments are less numerous in smaller worms. The filaments are united for the first 3 mm of their length by a thin basal web (B.W., Fig. 1A) running round the abfrontal face of the crown base, but distal to this they are entirely separate.