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Studies on the Animal Symbionts of the Gorgonian Coral, Leptogorgia Virgulata (Lamarck)

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Studies on the Animal Symbionts of the Gorgonian Coral, Leptogorgia Virgulata (Lamarck) STUDIES ON THE ANIMAL SYMBIONTS OF THE GORGONIAN CORAL, LEPTOGORGIA VIRGULATA (LAMARCK) WENDELL K. PATrON Ohio Wesleyan University, Delaware, Ohio 43015 ABSTRACT Colonies of Leptogorgia in the vicinity of Beaufort, North Carolina, occur most commonly in shades of yellow or orange and harbor several species of symbiotic animals whose adaptations are discussed. The snail Neosimnia uniplicata feeds on surface debris and material shed by the host and incorporates pigment from the host colony into its own shell. The gorgonian also harbors a small shrimp, Neopontonides beau/ortensis; a tissue-feeding nudibranch, Tritonia wellsi; a bivalve, Pteria colymbus; and two unidentified species of copepods. The barnacle Balanus galeatus settles preferentially on the decaying egg masses of Neosimnia. It grows down to the axis of the gorgonian and, following the disappearance of the egg mass, is surrounded by host tissue. INTRODUCTION The sea whip Leptogorgia virgulata (Lamarck) is a branching gorgonian or horny coral which grows to an overall length of about 60 centimeters and is quite common on hard, subtidal substrates in the vicinity of Beaufort, North Carolina. It harbors several quite specialized symbiotic animals. These are shown diagrammatically in Figure 1 and are the subject of this paper. A portion of this material has appeared in abstract (Patton, 1963). MATERIALS AND METHODS A total of about 200 colonies of Leptogorgia were collected over a period of several years from various sites in the Beaufort area and examined for symbionts. The chief collecting site was the rock jetty on Radio Island, a locality inside Beaufort Inlet subject to fairly strong currents. Collections were made from 1 to 10 feet below mean low tide and were made primarily during the summer months. To reduce the loss of symbionts during col- lection, each colony was placed in a hand net, then freed from the substrate and brought to the surface. The symbionts were placed on pieces of Leptogorgia in dishes of sea water and observed under the microscope. Long-term experiments and observations were hampered by the fact that colonies of Leptogorgia kept in running sea water in the laboratory usually decomposed within a week, perhaps due to the large amount of sediment in the water. Survival was 420 Bulletin of Marine Science [22(2) lem FIGURE 1. Diagrammatic drawing of a colony of Leptogorgia and its symbionts: 1, Neosimnia uniplicata with mantle withdrawn; 2, N. uniplicata with mantle extended; 3, egg mass of N. uniplicata; 4, Pteria colymbus; 5, Neopontonides beaufortensis; 6, Tritonia wellsi; 7, Balanus galeatus. better in aerated tanks of artificial sea water. Brine shrimp nauplii were added to these tanks at regular intervals, and, although the polyps were never seen fully extended, specimens seemed to be in good condition for the first several weeks and survived up to 3 months. 1972] Patton: Animal Symbionts of Gorgonian Coral 421 THE HOST Leptogorgia virgulata, like other gorgonians, is a colonial organism and is composed of a central axis of protein and an outer layer of tissue known as coenenchyme. The polyps extend out of the coenenchyme into the water, but can be withdrawn, leaving a pattern of holes which are darker than the surrounding tissue. The polyps of L. virgulata are white and translucent and lack the symbiotic zooxanthellae found in many other species of gorgonians. In whole colonies in the field and in pieces of freshly collected Leptogorgia in laboratory dishes, the polyps tended to be expanded under reduced illumination and contracted under bright light, although this relationship was by no means absolute. Embedded within the coenenchyme are small, colored, calcareous spicules. The color of Leptogorgia varies from specimen to specimen, but, with only rare exceptions, is constant throughout the length of a given colony. In the Beaufort area, specimens of L. virgulata occur most com- monly in shades of yellow and orange, while red, purple, and white colonies are rare. In Chesapeake Bay to the north and at Alligator Harbor, Florida, in the Gulf of Mexico, purple colonies are common. The central axis is thin at the tip of a branch, becoming progressively heavier toward the base, and finally forming a plate of axial material which attaches to the substrate and enlarges as the specimen grows. The axis is normally covered by coenenchyme, but, particularly at the base of large colonies, may be exposed and thus available as a site for settlement by fouling organisms. Throughout the period when colonies of Leptogorgia were maintained in tanks of aerated water, thin sheets of spicule-containing material were seen to be shed by the gorgonian. This did not seem to be associated with decomposition and appeared to be a natural process. S. A. Wainwright and G. R. Murdoch (personal communication) have observed a similar phenomenon in the gorgonian Pterogorgia citrina. Under both field and laboratory conditions, this species sheds transparent, noncellular organic membranes that have spicules stuck to them and are fairly well covered with diatoms and other microorganisms. THE SYMBIONTS Neosimnia uniplicata (Sowerby).-This is an elongate, thin-shelled, proso- branch gastropod and is the best known of the symbionts of Leptogorgia, having been reported by Osburn (1885) and several subsequent authors. It belongs to the Ovulidae, a fairly small family, all of whose members seem to live in association with cnidarians (Robertson, 1970). Several snails were usually found together on a host. In a related genus, Cyphoma, the occurrence of male-female pairs and intraspecific fighting behavior has been observed (Ghiselin & Wilson, 1966). This was not noticed in N. uniplicata. 422 Bulletin of Marine Science [22(2) With regard to feeding habits, specimens of Neosimnia are found moving up and down the branches of the colony with their oral area pressed against the surface. Occasionally an individual was seen to bite the tissue at the edge of the pore formed by a withdrawn polyp or to pull tissue from the exposed cut end of a branch. In the great majority of cases, however, there was no indication that living tissues were ingested, nor did microscopic examination of the gorgonian after a snail had passed reveal any sign of damage. However, both freshly collected and laboratory specimens of Neosimnia produce feces that contain gorgonian spicules. Although direct ingestion of living tissue of the host occasionally occurs, it is probable that material settling on the colony and the above-mentioned thin sheets of spicule-containing material shed by the host are major components of the diet. This would account for the presence of spicules in the feces and for the lack of damage to the host. When starved snails were placed on pieces of Leptogorgia that had been rubbed gently in carmine powder, the snails moved over the gorgonian and rapidly ingested the carmine and the host's mucus in which the carmine particles were entangled. N. uniplicata evidently has a more benign relationship with its host than do several other species of the family Ovulidae, which remove quite notice- able amounts of host tissue. Simnia spelta feeds on host coenenchyme and may denude portions of branches of the gorgonian Eunicella stricta (Theodor, 1967). A photograph (Berrill, 1966) shows two individuals of Cyphoma removing the coenenchyme from a specimen of Pseudopterogorgia and ex- posing the central axis. Simnia patula eats the flesh of the alcyonarian Alcyonium digitatum (Lebour, 1932), including contracted polyps (Fret- ter&Graham, 1962). R. Robertson (personal communication) has noticed scars on an alcyonarian caused by Calpurnus verrucosus. Females of N. uniplicata deposit gelatinous masses containing a variable number of egg capsules around the branches of Leptogorgia (Fig. 1). The hundreds of eggs in each mass are initially white, but become darker as de- velopment proceeds and hatch as brown-shelled veligers. The gelatinous mass remains in place for some time after hatching occurs and serves as a site for the attachment of algae and other organisms. Eventually, however, the mass breaks apart and drops off the branch. The host coenenchyme underneath the egg mass occasionally disintegrates, exposing the axis. It is much more common, however, for the tissue to be somewhat contracted but still appear healthy. This is quite unlike the situation in the gorgonian Eunicella stricta, where the egg masses of Simnia spelta usually cause necrosis of the underlying tissue. This exposes the central axis to settlement by fouling organisms. In some cases these organisms can withstand being overgrown and enclosed by the regenerating coenenchyme (Theodor, 1964, 1967). 1972] Patton: Animal Symbionts of Gorgonian Coral 423 TABLE 1 DISTRIBUTION OF SHELL LENGTH IN Two COLLECTIONS OF Neosimnia uniplicata Number of specimens Shell length (mm) June 6-7, 1961 August 16, 1961 3.0- 4.9 1 5.0- 6.9 5 7.0- 8.9 13 9.0-10.9 12 11.0-12.9 6 11 13.0-14.9 16 7 15.0-16.9 21 2 17.0-18.9 6 1 19.0-20.9 5 Since Leptogorgia was not collected during the winter months, the exact period of egg-laying by Neosimnia is not known. A single egg-containing mass was found in the earliest collection, March 29, and in the latest one, November 13. In early June, egg masses of all stages from freshly deposited to completely spent are common, but young snails are not yet found (Table 1). As the summer progresses, young snails become common, and the largest specimens evidently die off (Table 1). In studying the develop- ment of Simnia spelta, Thiriot-Qui6vreux (1967) found that larval life lasted 2 to 3 months, a timespan which seems plausible for Neosimnia uniplieata. Osburn (1885), Field (1949) and Perry & Schwengel (1955) have all noted that the color of the shells of N.
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