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<I>Sphaeroma Terebrans</I> BULLETIN OF MARINE SCIENCE, 76(1): 27–46, 2005 THE DISTRIBUTION AND ABUNDANCE OF SPHAEROMA TEREBRANS, A WOOD-BORING ISOPOD OF RED MANGROVE (RHIZOPHORA MANGLE) HABITAT WITHIN TAMPA BAY R. Allen Brooks and Susan S. Bell ABSTRACT This study was conducted to determine the distribution, abundance, and demog- raphy of a wood boring isopod, Sphaeroma terebrans Bate, 1866, within the prop roots of the red mangrove, Rhizophora mangle L., in eight sites within Tampa Bay, Florida. Sphaeroma terebrans in Tampa Bay displayed reproductive activity year- round and bay-wide synchrony in their density pattern. On average approximately 60% (range: 25%–86%) of the intertidal aerial roots surveyed were occupied by S. terebrans. Although infestation levels by S. terebrans in Tampa Bay were similar to that of more tropical regions, the distribution of S. terebrans was not continu- ous throughout the study sites. A substantially higher occurrence and density of S. terebrans was found in the northern compared to more southern study sites within the Bay. Additionally, some seemingly suitable areas of the bay (i.e., Pinellas Point, Skyway, Fort Desoto) were actually unoccupied on some dates. Although sites dif- fered in the frequency with which roots were attacked, the density of burrows and isopods in an occupied root was similar, with most attacked roots containing 3–5 burrows. The results of a transplantation experiment indicated that neither abiotic factors nor substrate quality limit the burrowing capabilities or survival of adult S. terebrans in the areas where they are absent. Instead, dispersal limitation, linked with differential juvenile survival, most likely controls isopod distribution and abundance within Tampa Bay. Sphaeroma terebrans (Bate, 1866), a wood boring isopod, is distributed worldwide in tropical mangroves (Estevez, 1978). Arguably, S. terebrans is not native to North and South America but was introduced from the Indo-Pacific when isopods bored into the hull of wooden shipping vessels (Carlton, 1994). Regionally, within North America, geographical surveys indicate that S. terebrans is distributed continuous- ly along both the east and west coast of Florida but, curiously, is not found in the Florida Keys (Conover and Reid, 1975; Rice et al., 1990). The isopod is found farther south, however, in the tropics of Central and South America (Ellison and Farnsworth, 1990). Local within-stand occupation by S. terebrans has been noted to be patchy with seemingly habitable areas unoccupied (Estevez, 1978). Abiotic differences in sa- linity, water temperature, dissolved oxygen, suspended solids, and flow have all been suggested to potentially influence the distribution of S. terebrans (Conover and Reid, 1975; Estevez, 1978; Barkati and Tirmizi, 1990). Ellison and Farnsworth (1990) and Ellison et al. (1996), also found that root fouling by sponges and colonial ascidians can impact isopod distribution. Sphaeroma terebrans exploits habitat created by mangrove prop roots and, in Florida, bores almost exclusively into unattached aerial roots of the red mangrove, Rhizophora mangle L. Once constructed, the burrow is used for 1) protection from both abiotic (exposure, desiccation) and biotic factors (Estevez, 1978); 2) filter-feed- ing activities (suspended sediment, algae, and bacteria; Rotramel, 1975; Rice et al., 1990); and 3) reproduction along with maternal care (Thiel, 1999). It is common to find inquilines (i.e., cohabiting amphipods, annelids, isopods) utilizing both occu- Bulletin of Marine Science 27 © 2005 Rosenstiel School of Marine and Atmospheric Science of the University of Miami 28 BULLETIN OF MARINE SCIENCE, VOL. 76, NO. 1, 2005 pied and unoccupied burrows of S. terebrans (Estevez, 1978). The distribution of S. terebrans within a mangrove stand is limited to intertidal regions that are flooded on a regular tidal cycle (Estevez, 1978) and abundance cycles of S. terebrans have been linked with seasonal fluctuations in salinity, temperature, water cover, and food lev- els (Estevez, 1978). An understanding of the factors that impact mangrove root occupation by S. ter- ebrans is important as the activity of this woodborer can have community-wide im- pacts. Isopod attack impacts the mangrove tree directly through root architectural changes (Simberloff et al., 1978; Ribbi, 1981), reduced root production, and increased root atrophy (Perry, 1988; Perry and Brusca, 1989; Ellison and Farnsworth, 1990). These changes to the root system not only alter support and nutrient provisioning for the tree but also may indirectly affect organisms that utilize the mangrove roots as either substratum (Sutherland, 1980; Rodriguez and Stoner, 1990; Bingham, 1992; Ellison and Farnsworth, 1992) or protective habitat (Primavera, 1997). Additionally, S. terebrans may have important economic impacts when in high abundance due to considerable damage to maritime structures (Rice et al., 1990; Cragg et al., 1999). This study expands upon the work of Estevez (1978) and addresses two main ques- tions regarding S. terebrans inhabiting prop roots of R. mangle within Tampa Bay: 1) Is the current distribution of S. terebrans continuous within the mangrove habitat of the Tampa Bay region, and if not, then what factors might explain the disjunct distri- bution?; and 2) Do abundances of S. terebrans differ among the mangrove stands in which they are found in Tampa Bay and, if so, can the differences be accounted for by the demographic features of fecundity, juvenile recruitment, or sex ratios? MATERIALS AND METHODS SITE SELECTION.—Eight sites around Tampa Bay (Fig. 1) were selected for study based upon their geographic range from the northern to southernmost part of the bay. All of the sites con- tained areas of continuous mangrove coverage dominated by red mangroves. Based upon abi- otic information obtained from the Hillsborough County Ecological Protection Commission monitoring stations, northern sites on average experience a lower turbidity and salinity level but similar pH, dissolved oxygen, and water temperature compared to the more southern sites within the bay. Within each of the eight locations ten sampling points were haphazardly established along the seaward edge in the intertidal region. Each sampling point consisted of a single R. mangle tree that was tagged with forestry tape. PRESENCE/ABSENCE CENSUS.—Transect samples were utilized to determine the frequency of root attack by S. terebrans at all eight field sites using a method similar to Ellison and Farn- sworth (1990). Roots were sampled to determine isopod density in October 1999, February 2000, and July 2000 at all sites. These specific months were selected as they encompassed times of both high and low isopod abundance (Estevez, 1978). All sites were sampled within 2 wks of each other. Eight transects, 25 m in length, were haphazardly placed parallel to the seaward edge of the mangrove habitat in each study site. Along each transect, 30 random points were selected for tallying of isopod presence/absence. At each random point along the transect the closest unattached aerial roots with one tip was selected for evaluation. An unattached root is a prop root originating from the bole, trunk, and other aboveground roots, which grows down through the water column and eventually attaches to the substratum (Gill and Tomlinson, 1977). Selected roots were located within 1 m of the seaward edge and sub- merged regularly at high tide. Isopod presence was noted if there was visible sign of burrow- ing activity along the root (≥ 1 burrow). BROOKS AND BELL: DISTRIBUTION AND ABUNDANCE OF SPHAEROMA TEREBRANS 29 Figure 1. The eight study sites around Tampa Bay. Abbreviations are as follows: UTB = Upper Tampa Bay, 4th = Fourth Street, WI = Weedon Island, CRB = Cockroach Bay, PP = Pinellas Point, SKY = Skyway, FD = Fort Desoto, and AM = Anna Maria Island. The distance between sites UTB and AM is 51 km. Map of Tampa Bay courtesy of the Florida Marine Research Insti- tute, Florida Fish and Wildlife Conservation Commission. ROOT SAMPLING PROCEDURE.—Root sampling was conducted on the same schedule as the transect surveys. Roots were not sampled from Pinellas Point and Skyway because of extremely low isopod abundance within those sites. Similarly, roots were only sampled from Fort Desoto in February and July. Unattached aerial roots located within 1 m of the seaward root edge were destructively sampled for isopod density estimation. Additionally, only roots which displayed signs of burrowing by S. terebrans (≥ 1 burrow) were sampled. Chosen roots contained only one root tip as previous studies suggest that isopods prefer root tips and there- fore the presence of multiple root tips might bias density estimates studies (e.g., Perry and Brusca, 1989; Brooks and Bell, 2001a). The five closest attacked roots (which met the above stated criteria) to each sampling point were taken. In the laboratory, all burrows were noted as to their spatial location along the root. Bur- rows were then excavated to determine the presence, size, sex, and reproductive status of S. terebrans colonizers. Individuals were pressed flat under a dissecting scope and the length from the tip of the head to the tip of the pleotelson was recorded (Thiel, 1999). Juveniles were defined as individuals < 5.6 mm and possessed no sign of sexually dimorphic features, in ac- cordance with Estevez (1978) and Venkatakrishnana and Nair (1973). Males were identified by the presence of penes (Venkatakrishnana and Nair, 1973). Adult females were classified as brooding if there were either eggs or embryos present within the brood chamber. Maternal care in which juveniles are clustered at the terminal end of a family burrow (sensu Thiel, 1999) 30 BULLETIN OF MARINE SCIENCE, VOL. 76, NO. 1, 2005 was recorded separately. Additionally, any root that contained an unoccupied burrow inhab- ited by an inquiline isopod was recorded. HABITAT AVAILABILITY.—The density of unattached aerial roots at each study site was de- termined using a 1 m2 collapsible quadrat placed within the prop roots adjacent to each sam- pling point. The front edge of the quadrat was aligned parallel to the forest edge, which was defined in this study as the furthest extending prop root at the land/water ecotone.
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