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<I>Diopatra Cuprea</I>

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<I>Diopatra Cuprea</I> BULLETIN OF MARINE SCIENCE, 40(1): 11-21, 1987 ROLE OF DIOPATRA CUPREA BOSC (POLYCHAETA: ONUPHIDAE) TUBES IN STRUCTURING A SUBTROPICAL INFAUNAL COMMUNITY Suzanne M. Ban and Walter G. Nelson ABSTRACT An a priori hypothesis predicted that in the vicinity of aggregated Diopatra cuprea tubes an enhanced infaunal density and species richness would be found, resulting from a biological refuge effect of the tubes. To test this hypothesis, cores were taken over a 5-month period in both vegetated, Halodule wrighti! Aschers. beds, and unvegetated areas of a site in the Indian River lagoon, Florida. An inner, 0.01 m2, frame was placed to enclose densities of 0, I, or 4 D. cuprea tubes, while an outer concentric, 0.02 m2, frame was placed so that it enclosed the smaller frame, plus a surrounding area lacking in D. cuprea tubes. The presence of D. cuprea tubes was found to have no consistent significant effect on the abundance and number of infaunal species found in either the vegetated or unvegetated areas. Laboratory experiments employing a benthic predator, Callinectes, were carried out in order to determine whether D. cuprea tubes andlor H. wrightii rhizome mats actually constitute a barrier to predation. Significantly higher survivorship of the bivalve Mulinia lateralis Say, used as prey, was found in laboratory treatments containing 10 tubes per 0.01 m2 versus treatments containing 4 or a tubes per 0.01 m2. Highest survivorship of bivalves was found in treatments containing a H. wrightii rhizome mat; tubes placed within the mat did not enhance clam survivorship. The discrepancy between the findings of this study, and previous studies on the refuge effect of D. cuprea tubes which found evidence to support the refuge hypothesis, indicates that there may be a critical lower limit of tube density that is needed to establish an effective refuge. This density was not found at the Indian River study site and may account for the lack of an observed refuge effect in the field data. Natural disturbance can be important as a factor influencing the community structure of the marine benthos (Woodin, 1978; Thistle, 1980; 1981; Miller, 1982). Biological perturbation by motile predators (e.g., decapod crustaceans, Virnstein, 1977; Woodin, 1978; 1981; Holland et a1., 1980; Nelson, 1981; bottom- feeding fishes, Arntz, 1977; Virnstein, 1977; shorebirds, Goss-Custard, 1977; Luckenbach, 1984) can be a significant source of mortality to the infaunal benthos. Physical structures that reduce the susceptibility ofthe infauna to mortality caused by either physical or biological disturbance have been termed refuges (Woodin, 1978). Several biologically generated refuge structures have received recent study. The density and diversity of infaunal organisms have been found to be increased in association with biogenic refuges such as seagrass beds (Young et a1., 1976; Orth, 1977; Reise, 1977; Nelson, 1981; Peterson, 1982; Virnstein et a1., 1983), oyster shells (Dauer et a1., 1982), and worm tubes (Woodin, 1978; 1981; Wilson, 1979). Brenchley (1982) found that two kinds of biogenic structures, roots of seagrass and tubes of invertebrat~s, are able to deter the mobility of a number of burrowing predators. The combined effect of roots and tubes in the substrate has the greatest impact on predator mobility. The tube structures of the onuphid polychaete Diopatra cuprea have been found to be one such biogenic refuge. Woodin (1978; 1981) found elevated infaunal density and diversity around aggregated D. cuprea tubes in a Virginia mud flat. In addition, Luckenbach (1984) found that D. cuprea tubes appear to provide an effective refuge for macrofauna from shorebird predation. However, in a recent II 12 BULLETIN OF MARINE SCIENCE, VOL. 40, NO.1, 1987 study, Belland Woodin (1984) found the spatial patterns ofmeiofaunaand selected macrofauna in relation to tubes to be highly variable. The discrepancy between the findings of Bell and Woodin (1984) and earlier reports of increased macrofaunal density in D. cuprea rich areas (Woodin, 1978; 1981) suggests that a number of unanswered questions remain. Most previous work on the refuge effect of D. cuprea tubes has been conducted in temperate areas. First, it is important to investigate whether results found in these temperate zone studies can be extrapolated to a functionally different subtropical system where small decapod predators are more numerous (Virnstein, 1977; 1978) and D. cuprea densities are lower. In addition, an understanding of the relative im- portance of the refuge effect of D. cuprea tubes within H. wrightii beds is needed since vegetated areas constitute a large part of subtropical subtidal systems. Fi- nally, although Woodin (1978; 1981) suggests that disturbance to the infauna by Callinectes sapidus Rathbun is reduced by the presence of D. cuprea tubes, she does not document their specific behavior around the tubes. The behavior of Callinectes in the vicinity of aggregated D. cuprea tubes must be examined in order to determine whether in fact tubes constitute an effective refuge from Cal- linectes predation. The present study seeks to examine these three questions. METHODS AND MATERIALS Field Study. -Fieldwork was conducted at a site 1.5 km south of the Sebastian Inlet in the Indian River lagoon, Florida (80029'W, 27°50'N). The study site was located in the shallow subtidal zone along a mangrove dominated shoreline. The bottom closest inshore consisted of interspersed vegetated and unvegetated patches, grading into larger Halodule wrightii grassbeds in the offshore direction. An offshore sandbar separated H. wrightii grassbeds from those dominated by Syringodium filiforme. Maximum depth inshore of the sandbar was approximately I m. A detailed description of the study site can be found in Ban (1985). To examine the association between the presence (and/or abundance) of D. cuprea tubes and the abundance and diversity of the remainder of the infauna, sediment cores were taken from both vegetated H. wrightii beds and unvegetated sandy areas following the methods of Woodin (1978). Sampling was done over a 5-month period, from May to September 1983. Observations at the site indicated highest D. cuprea population densities were found during this period. Sampling was dis- continued in October 1983 due to the low density of D. cuprea. Sampling employed the use of two aluminum frames, 0.01 m2 x 14 cm deep and 0.02 m2 x 14 cm deep. The smaller frame.was oriented so that it enclosed an area with 0, 1, or 4 D. cuprea tubes. The maximum of 4 tubes was chosen because this is the largest number of worm tubes that could be consistently found together in 0.0 I m2• The outer 0.02-m2 frame was placed so that it enclosed the smaller frame plus the surrounding area containing 0 D. cuprea. Thus, two concentric 0.01 m2 x 14-cm-deep samples were obtained, the inner containing a known density of worm tubes, the outer containing no D. cuprea. Tube caps and seagrass blades were cut off at the sediment surface prior to the removal of the core in order to minimize contamination by epifaunal organisms. Sampling was done at low tide, in water depths of between 10 and 30 cm, to facilitate the search for the tubes and the removal of the cores. The tidal range in this area is small, less than 50 cm, and subject to weather conditions (G. Swain, pers. comm.). Three replicates of each D. cuprea density were taken each month. Upon removal from the substrate with a shovel, cores were separated and washed on 0.5-mm Nitex mesh squares to remove sediment. Samples were immediately fixed in 7% Formalin. At least 48 h later, the preserved samples were again washed through a 0.5-mm screen and stored in a 70% alcohol- rose bengal solution. These samples were sorted, identified, and only whole animals and heads were enumerated. Samples were tested by three-way analysis of variance (ANOVA) with replication to analyze the response of macrofauna I abundance and species richness to the main effects of time, D. cuprea density, and vegetation presence or absence. Prior to computation of the ANOV A statistics, the homogeneity of variances was checked by the Bartlett test (Sokal and Rohlf, 1981) and a 10glO(x+ I) transformation was employed where needed. Laboratory Experiments. - These were conducted in glass aquaria each divided into three sections, designed to allow direct observation and determination of the effect of D. cuprea tubes on the foraging techniques of Callinectes. Juvenile Callinectes (approximately 8 cm in carapace width) were collected in the early spring from the Sebastian sampling site. The crabs were only identified to the level of BAN AND NELSON: ROLE OF DIOPATRA CUPREA TUBES IN AN INFAUNAL COMMUNITY 13 genus. The dominant species in this area is Callinectes sapidus, but C. ornatus and C. similis are also present (Gore et aI., 1981). Mulinia latera lis of the size range 10-20 mm were collected from the Indian River and used as a food source. Sediment from the field study site was allowed to become azoic by air drying for at least 30 days prior to the commencement of the experiment, and was used as a sediment base in the aquaria. Experiment I tested the null hypothesis that higher tube densities are more effective in deterring crab foraging. Three treatments, each replicated three times were used: I) a tube density of 4/0.0 I m2, the maximum consistently found at the Indian River study site; 2) a tube density of 10/0.01 m2, the maximum found by Woodin (1978) at Tom's Cove, Virginia; and 3) zero tubes, to act as a control. Large D. cuprea tubes collected from the field study site were pushed into the sediment and reinforced from the inside with toothpicks.
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