Feeding Habits of <I>Oreaster Reticulatus</I>
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BULLETIN OF MARINE SCIENCE, 32(2): 504-510, 1982 FEEDING HABITS OF OREASTER RETICULATUS (ECHINODERMATA: ASTEROIDEA) R. E. Scheibling ABSTRACT In situ feeding observations among eight populations in various grassbed and sand-bottom habitats in the Grenadines and off St. Croix. U.S. Virgin Islands indicate that Oreaster reticulatus is primarily an omnivorous. microphagous substratum grazer. It extrudes its extensive cardiac stomach upon seagrass. sand, and/or algal substrates and ingests associated micro-organisms and particulate detritus. O. reticulatus also functions as an opportunistic predator and scavenger of any available slow-moving, sessile or moribund macrofauna and is occasionally cannibalistic. The feeding habits of O. reticulatus may be exemplary of tropical, shallow-water asteroids in general. Oreaster reticulatus (Linnaeus) is an abundant and conspicuous inhabitant of seagrass and sand bottoms in the Caribbean (Scheibling, 1980a,b). However, published information on its feeding biology is limited. A few incidental obser- vations and circumstantial evidence have led to divergent conclusions: O. reti- culatus typically digests microscopic organic material upon the substratum but occasionally feeds on sponges (Thomas, 1960); it preys upon the urchin Meoma ventricosa (Kier and Grant, 1965); and, based on anatomical specialization of the gut, it functions as a suspension feeder (Anderson, 1978). Anderson (1978) has expressed the need that "studies soon be made on the feeding biology of Oreaster reticulatus, to supplement incidental observations, and to determine the validity of conclusions that can now be only inferred from considerations of the anatomy of the digestive system." This study surveys the feeding habits of Oreaster reticulatus in situ in several populations from various habitats in the Carribbean. Microphagous substratum grazing and macrophagous predation and scavenging are examined as alternate feeding modes in relation to anatomical specializations of the digestive tract and food availability. MATERIALS AND METHODS In situ observations of feeding in Oreaster reticulatus were conducted among populations in grassbed habitats (Hillsborough Bay and L'Esterre Bay, Carriacou Island, and Chatham Bay, Union Island, Grenadines) and in sand-bottom habitats (Watering Bay and Manchioneal Bay, Carriacou Island, and Buck Island Channel, Coakley Bay and Horseshoe Patch, St. Croix, U.S. Virgin Islands) by overturning individuals and noting the type of substratum or macroscopic prey encompassed by the broadly extended stomach. O. reticu/atus were sampled systematically in transect surveys (Schei- bling, 1980a,b) or as encountered in unidirectional traverses of a study site. Ingested particulate organic matter was collected from the stomachs of individual Oreaster reti- cu/atus feeding upon sand substrata in Buck Island Channel and Horseshoe Patch. The asteroids were overturned rapidly and the nozzle of a 50-ml syringe was inserted into the closing mouth. (Disturbance evokes retraction of the stomach and closure of the mouth and ambulacral grooves within 3-5 sec.) With the syringe clamped by the mouth, a 10-ml sample of seawater, entrapped within the lobes of the cardiac stomach during retraction, was withdrawn. Ten samples (different individuals) were pooled for each study site and centrifuged for 15 min in a Sorvol high-speed cen- trifuge; the particulate residue was examined microscopically. The density of sea urchin prey of Oreaster reticu/atus was measured concurrently with that of the asteroid. In Chatham Bay, Tripneustes ventricosus were enumerated in 32, 25-m2 quadrats placed at 25-m intervals along 8, IOQ-m long transects (Scheibling, 1980a). In Horseshoe Patch, Meoma ventricosa were enumerated in 28, 10-m diam. circular census stations (Scheibling, I980b). In 504 SCHE1BLlNG: FEEDING HABITS OF OREASTER RETICULATUS 505 Hillsborough Bay, a 16-m2 wire mesh enclosure was set in the Ha/odu/e wrightii zone (Scheibling, 1980a) and stocked with 20 O. reticu/atus and 20 T. ventricosus. The predation of T. ventricosus by O. reticu/atus in the enclosure was monitored for 3 days. Asteroids that escaped and urchins that were preyed upon were replaced daily. RESULTS The summary of 10,328 in situ observations of feeding in Oreaster reticulatus from three populations in grassbed habitats and five populations in sand-bottom habitats is given in Table I. Feeding is divided into two basic types: (I) substratum grazing, in which the cardiac stomach is everted upon the substratum and func- tions as a microphagous feeding organ, and (2) predation and scavenging, in which the stomach envelops macrofaunal prey or carrion. Substratum grazing is sub- divided according to the predominant substratum type (e.g., sand, seagrass, al- gae). Microphagous substratum grazing is the primary feeding type of O. reti- culatus in nature, accounting for 99-100% of all feeding observations in 7 of the 8 populations studied and 90% in the remaining one (Chatham Bay). Substratum Grazing.-In grassbed habitats, Oreaster reticulatus fed prefer- entially upon Halodule wrightii (Table 1 and Scheibling, 1980a). Grassblades were clustered beneath the disc by the tube feet and enveloped by the everted cardiac stomach. Degradation of the blades was not discernible, indicating that O. reticulatus was probably digesting epiphytic micro-organisms, filamentous algae, and particulate detritus associated with H. wrightii and not the seagrass per se. Dense mats of filamentous green algae (Chaetomorpha, Enteromorpha) were intensively grazed by O. reticulatus during periodic blooms (2-3 mo inter- vals). These mats covered from 50-75% of the bottom and harbored abundant micro-organisms and detritus. O. reticulatus fed upon bare carbonate sand in pockets amid the beds of Halodule wrightii, in an inshore sandy zone (Hillsbor- ough Bay and L'Esterre Bay), and in patches of sparse Thalassia testudinum (Chatham Bay). Infrequently, O. reticulatus everted its stomach upon clusters of dead, detached blades of T. tesludinum ("Other Substratum Grazing" in Table I) that had collected in the sand pockets, probably to digest films of bacteria and other micro-organisms on the surface of the litter. In sand-bottom habitats, Oreasler reticulatus fed almost exclusively upon the bare sand substratum (Table 1). The carbonate surface sediments were raked by the tube feet (in the same manner as grassblades) to form a low mound beneath the disc upon which the cardiac stomach was everted (Scheibling, 1980d). Stomachs of feeding individuals in Buck Island Channel and Horseshoe Patch contained an array of particulate organic matter including: numerous dia- toms; dinoflagellates, foraminifera and other protozoans; filaments of green and bluegreen algae; meiofaunal crustaceans (e.g., harpactacoid copepods, ostra- cods, crab larvae), polychaetes, nematodes; and bacterio-detrital aggregates. O. reticulatus was infrequently observed with its stomach everted upon seagrass (Syringodiumfiliforme) or red macroalgae (Lophocladia trichoclados, Wrangelia biscuspidata, Laurencia corallopsis, Wurdemania miniata) which occurred spo- radically in sand-bottom habitats. The red algae were discolored (orange or red) where in contact with the stomach. In Watering Bay, O. reticulatus was infre- quently observed with its stomach everted upon coral rubble ("Other Substratum Grazing" in Table 1) apparently digesting organic films and/or encrusting coralline algae on the rubble surface. Predation and Scavenging.-Oreaster reticulatus infrequently preyed upon ses- sile or slow-moving macrofauna. In the grassbed populations, the sea urchin 506 BULLETIN OF MARINE SCIENCE. VOL. 32. NO.2. 1982 Table l. Tbe feeding habits of Oreaster reticulatus in populations from (a) grass bed and (b) sand- bottom habitats: percentage frequencies of substratum grazing on various substratum types, and of predation/scavenging Percentage of Feeding Observations Pre- Number of Substratum Grazing dation! Feeding Scaveng- Populations Observations Sand Grass Algae Other Total ing (a) Grassbed Habitats Hillsborough Bay 3,346 7.7 74.4 ]7.2 0.1 99.4 0.6 L'Esterre Bay 374 ]5.4 74.5 9.3 0.4 99.6 0.4 Chatham Bay 611 19.8 48.5 21.9 0.] 90.3 9.7 (b) Sand-Bottom Habitats Buck I. Channel 1,833 98.4 0.3 1.2 0 99.9 0.1 Coakley Bay 2,355 95.1 0 4.9 0 100.0 0 Horseshoe Patch 1,162 95.2 2.2 1.9 0 99.3 0.7 Watering Bay 568 %.2 0 0 2.6 98.8 1.2 Manchioneal Bay 79 100.0 0 0 0 100.0 0 Tripneustes ventricosus was a predominant prey item. Predation of T. ventricosus by O. reticulatus was rare in Hillsborough Bay and L'Esterre Bay (7 out of a total of 3,720 feeding observations) due to the paucity of the urchin on beds of Halodule wrightii inhabited by the asteroid. When offered T. ventricosus in an enclosure in the H. wrightii zone, 10-20 O. reticulatus consumed an average of six urchins/day (range: 4-9) over a 3-day period. Predation of T. ventricosus was more frequent in Chatham Bay (43 out of 611 feeding observations) and increased with urchin density within various substratum types (Table 2). The density of O. reticulatus, however, was greatest in beds of H. wrightii where T. ventricosus was least abundant. Oreaster reticulatus attacked Tripneustes ventricosus within a radius of 10-30 cm. The tip of the leading ray was curled upwards, and the terminal tentacles were extended and active as the asteroid moved towards its potential prey. Con- tact by the ray tip evoked vigorous spine activity, projection of globiferous ped- icellaria and a flight response in T. ventricosus. The pedicellaria affixed them- selves to the epidermis and tube feet of the ray tip, eliciting its momentary retraction, but pursuit followed. Chases lasting 5-10 min were observed; in each instance the faster moving urchin escaped O. reticulatus. T. ventricosus occa- sionally initiated flight prior to contact by an attacking asteroid. When feeding upon Tripneustes ventricosus, Oreaster reticulatus mounted and overlaid the urchin, enveloping the test with the fully everted cardiac stomach. Once the epidermis was digested, the stomach was inserted into the test through the periproct to digest internal tissue and gut contents of the urchin.