ENVIRONMENTAL IMPACT Sop RESEARCH PROGRAM df US Army Corps of Engineers TECHNICAL REPORT EL-84-4 y } Me 3 ECOLOGICAL EFFECTS OF RUBBLE WEIR JETTY CONSTRUCTION AT MURRELLS INLET, SOUTH CAROLINA VOLUME |: COLONIZATION AND COMMUNITY DEVELOPMENT ON NEW JETTIES by Robert F. Van Dolah, David M. Knott, Dale R. Calder South Carolina Wildlife and Marine Resources Department Marine Resources Research Institute Charleston, S. C. 29412 April 1984 Final Report Approved For Public Release; Distribution Unlimited Prepared for Office, Chief of Engineers, U. S. Army Washington, D. C. 20314 Under EIRP Work Unit 31532 Monitored by Coastal Engineering Research Center U. S. Army Engineer Waterways Experiment Station P. O. Box 631, Vicksburg, Miss. 39180 Destroy this report when no longer needed. Do not return it to the originator. The findings in this report are not to be construed as an official Department of the Army position unless so designated. by other authorized documents. The contents of this report are not to be used for advertising, publication, or promotional purposes. Citation of trade names does not constitute an official endorsement or appraval of the use of such commercial products. 30110001156 anoaraphic Institution Woods Hale Oceanographic In SECURITY CLASSIFICATION OF THIS PAGE (When Data Entered) =i READ INSTRUCTIONS REPORT DOCUMENTATION PAGE BEFORE COMPLETING FORM 1. REPORT NUMBER 2. GOVT ACCESSION NO.) 3. RECIPIENT’S CATALOG NUMBER Technical Report EL-84-4 4. TITLE (and Subtitle) 5. TYPE OF REPORT & PERIOD COVERED ECOLOGICAL EFFECTS OF RUBBLE WEIR JETTY CONSTRUCTION AT MURRELLS INLET, SOUTH CAROLINA; VOLUME I: COLONIZATION AND COMMUNITY DEVELOPMENT ON NEW JETTIES 7. AUTHOR(s) Robert F. Van Dolah David M. Knott Dale R. Calder 9. PERFORMING ORGANIZATION NAME AND ADDRESS Marine Resources Research Institute South Carolina Wildlife and Marine Resources Dept. Charleston, S. C. 29412 11. CONTROLLING OFFICE NAME AND ADDRESS Office, Chief of Engineers, U. S. Army Washington, D. C. 20314 Final report 6. PERFORMING ORG. REPORT NUMBER 8. CONTRACT OR GRANT NUMBER(e) 10. PROGRAM ELEMENT, PROJECT, TASK AREA & WORK UNIT NUMBERS Environmental Impact Research Program Work Unit 31532 12. REPORT DATE April 1984 13. NUMBER OF PAGES 138 15. SECURITY CLASS. (of this report) 14. MONITORING AGENCY NAME & ADDRESS(/f different from Controlling Office) Unclassified U. S. Army Engineer Waterways Experiment Station Coastal Engineering Research Center P. O. Box 631, Vicksburg, Miss. 39180 16. DISTRIBUTION STATEMENT (of this Report) DECL ASSIFICATION/ DOWNGRADING SCHEDULE 15a. Approved for public release; distribution unlimited. 17. DISTRIBUTION STATEMENT (of the abstract entered in Block 20, if different from Report) 18. SUPPLEMENTARY NOTES Available from National Technical Information Service, 5285 Port Royal Road, Springfield, Va. 22161. 19. KEY WORDS (Continue on reverse side if necessary and identify by block number) Colonization Jetties Fishes Motile macroinvertebrates Fouling community Murrells Inlet, South Carolina 20. ABSTRACT (Continue em reverse side ff meceseary and identify by block number) Quarrystone jetties constructed at Murrells Inlet, South Carolina, were studied over a 4-year period to evaluate community development patterns of biota colonizing the rocks. Sessile macroinvertebrates and algae were quantitatively assessed using line-transect and photographed-quadrat censusing techniques. Motile epifauna were also quantitatively sampled using a suction device, and fishes were qualitatively assessed using gill nets, hook and line, traps, seine net, and through visual observations while scuba diving. FORD DD . jan 7a 1473 EDITION OF 1 wov 65 1S OBSOLETE Unclassified SECURITY CLASSIFICATION OF THIS PASE (When Data Entered) Unclassified SECURITY CLASSIFICATION OF THIS PAGE(When Data Entered) 20. ABSTRACT (Concluded). The results documented that both jetties were rapidly colonized by sessile and motile biota. Within 1 year after construction, faunal and floral coverage of the rocks was equivalent to subsequent sampling periods, as were estimates of species diversity and abundance. Distinct vertical zonation of sessile biota was also observed within 1 year, with distribution patterns generally remaining similar throughout the study period. Vertical gradients in the distribution of motile fauna were less apparent, although some differences were noted intertidally versus subtidally. Community composition, on the other hand, changed both seasonally and yearly. Community structure appeared to change less over time in intertidal areas than in subtidal areas, where marked changes in dominant sessile taxa were observed between sampling periods. No stable or "climax" jetty community was apparent subtidally after 3 to 4 years, and other studies suggest that such a community is not likely to occur. Fish found around the jetties were abundant and included several recrea- tionally important species. Stomach content analysis indicated that the jetty biota was an important food resource for several fishes. In addition, at least one species, black sea bass, was using the rocks as a nursery area. Unclassified SECURITY CLASSIFICATION OF THIS PAGE(When Data Entered) PREFACE This report was sponsored by the Office, Chief of Engineers (OCE), U. S. Army, as a part of the Environmental Impact Research Program (EIRP) Work Unit 31532 entitled Ecological Effects of Rubble Structures, which was assigned to the U. S. Army Coastal Engineering Research Center (CERC). The Center, originally located at Fort Belvoir, Va., moved to the U. S. Army Engineer Waterways Experiment Station (WES), Vicksburg, Miss., on 1 July 1983. The Technical Monitors for the study were Dr. John Bushman and Mr. Earl Eiker of OCE and Mr. Dave Mathis, Water Resources Support Center. The study and preparation of a draft final report were accomplished during the time period September 1977 to May 1983. The report was prepared by Dr. Robert F. Van Dolah, Mr. David M. Knott, and Dr. Dale R. Calder through the Marine Resources Research Institute of the South Carolina Wildlife and Marine Resources Department. Dr. Calder is currently at the Royal Ontario Museum. The authors are very grateful to Mr. Arthur K. Hurme and Mr. E. J. Pullen of the CERC for their role in initiating this investigation, and for their support and encouragement throughout the study. We wish to thank Magdalene Maclin, Beth Roland and George Steele for their considerable efforts on this project, both in the field and laboratory. Other individuals who frequently assisted us in the field included Mary Jo Clise, Stan Hales, Priscilla Hinde, Terry Hodges and Caroline O'Rourke. Particular thanks are due to Dr. Reid Wiseman, who identified all of the algae found on the jetties, and to Dr. George Sedberry, who identified and analyzed the contents of fish stomachs. Finally, we wish to thank Nancy Beaumont who typed the various drafts of this report, and Karen Swanson who drafted all the figures. Mr. Hurme was the CERC Technical Advisor for the contract under the general supervision of Mr. Pullen, Chief, CERC Coastal Ecology Branch, and Mr. R. P. Savage, Chief, CERC Research Division. Dr. Roger T. Saucier, WES, was the Program Manager of EIRP. Technical Director of CERC at Fort Belvoir during the study and preparation of the draft final report was Dr. Robert W. Whalin. Commander and Director of WES during preparation of the final report was COL Tilford C. Creel, CE; Technical Director was Mr. F. R. Brown. This report should be cited as follows: Van) Dolah,, Re RY, Knott, D.) Mog and Calder, DeyR. 19845 kcoilloga call! Effects of Rubble Weir Jetty Construction at Murrells Inlet, South Carolinas; Volume I: Colonization and Community Development on New Jetties,'" Technical Report EL-84-4, prepared by Marine Resources Research Institute, Charleston, S. C., for Coastal Engineering Research Center, WES, Vicksburg, Miss. PREFACE LIST OF FIGURES ......ccccccccsscscces Jo0DD DDO DUD DOO OOD OOO DOD ONC HOOD DODD0S LEST OF TABLES 2.2... ccc ccc cecccscccescces SokohehleNeKeleNelelolenedeNoReleleeKeleleier le) elelelicleie Iho ILS IEILIE G IV. VII. INTRODUCTION .....--ceeese0 SOO ODDDD DD ODODODOOO ODDO O ODN DN 5000000000 DESCRIPTION OF THE STUDY AREA ....cccecsecrcccseccscccces 60000000000 MATERIALS AND METHODS 1. Station Characteristics and Sampling IWAN Gooodgao00gq00d000000 2. Sampling Dates 2.0.0. 35s s00- 5600600900000000000000006 90000000000 3. Biological Sampling Methods .......cccsecscccccrscererrescccces 4. Hydrographic Sampling .......-..0-- Sa0cooDO0GCDDDDDNN 60000000000 DD AleawAnauliy/Si'S mre reel etelevens 900000000 000000000 600000 op0000 0000000000 RESULTS AND DISCUSSION .......... 600000000 S600 0000000000000 ei eronederonene 1. Hydrographic Conditions ..........ceecececerrcccccccecs 50000000 2. Jetty Community Development ......... 360000 60000000000000000006 SUMMARY AND CONCLUSIONS .........2-- 300006 900000 s00000000000000000006 LITERATURE CITED ....... 660000 90000000006000 g00000000 SoC OUDOo0600 9 APPENDICES A. Percent cover of sessile macrofauna and flora estimated from line transects (75 pts.) at north jetty stations ....... B. Percent cover of sessile macrofauna and flora estimated from photographic quadrats (100 cm2) at north jetty SIEGIEMLOMS ooooocc og 0b DbOC DO DDKDO DODD ADDDDONDS o000000000 0090000000 C. Percent cover of sessile macrofauna and flora estimated from line transects (75 pts.) at south jetty stations ........ D. Percent cover of sessile macrofauna and flora estimated from photographic quadrats (150 cm2) at south jetty stations.. E. Line-transect estimates of total biota cover on rocks at the north and south jetty stations ...........ccccceoes 50000000 F. Ranked abundance of motile macroinvertebrates collected by slurp gun from.north jetty stations .............ceccecceccces G. Ranked abundance of motile macroinvertebrates collected by Siliusspe cunt rommESouthunie Etyprsica tO Sirepeleleleporsieheneierehelolokerenstelenoheleleleyo H. Estimates of species number, abundance, and diversity of TABLE OF CONTENTS ececececeec ere eo oer ece ee eee eee eee eo eee eee eee oe ee eee eT eee Oe Te Eo eT ee ee eo oO motile epifauna collected in suction samples at north and SOUEN JED, STARLOMS sooncte 00000 odd Dnd DOD DD DD NDDDNDDOODDOODNDNO 65 Al Bl Cl D1 EL Fl Gl H1 iLike A ESS 14. 1L3)e LIST OF FIGURES Page Map showing Murrells Inlet jetties and station locations ...... 9 Line-transect and photograph estimates of sessile biota cover atadiiterentmlievelisitok ithe north y et tyaieserselersretereisrrerctsicilienoron mre © Line-transect and photograph estimates of sessile biota cover atditherentleveilsvofmehes Souchiilettiyamorelishelelercilctelclereleneletcnepenctcnore Total number of sessile taxa observed at intertidal and subtidal levels of north jetty stations during line-transect census .... Total number of sessile taxa observed at intertidal and subtidal levels of south jetty stations during line-transect census .... Line-transect estimates of mean percent cover for the different sessile taxa found on the north jetty rocks ..........-...e-2ee0- Photographic estimates of mean percent cover for the different sessile taxa found on the north jetty rocks ..........-..-.e2--. Normal cluster dendrogram of north jetty line-transect data indicating station groups formed using the Bray-—Curtis Sahubllleyeliay ColenmeslerlEiMs oooqgno0 DDO Gdo0DK OGD DDOUDOGOObOUOOdOONDOND Line-transect estimates of mean percent cover for the different sessile taxa found on the intertidal south jetty rocks ........ Photographic estimates of mean percent cover for the different sessile taxa found on the intertidal south jetty rocks ........ Line-transect and photographic estimates of the mean percent cover for the different sessile taxa found at the -1.0m subtidal level on the protected side of the south jetty ...... : Normal cluster analysis of south jetty line-transect data indicating station groups formed using the Bray-Curtis phil healcizs (Kol meslOnkanye Mn nicgooddooOG OC epb00000000560600000000000 Vertical distribution of the 20 most abundant sessile species obsexvedjatunorth jetty stat lonspesceee oe oe eee Vertical distribution of the 20 most abundant sessile species obsenvedgatnsouthmjiettysstatilonsmmeeceeer eee Cena Linear regression of the abundance and number of species of motile epifauna at each north jetty station as a function OE tidal felievat dom: ci. 6 Mtereiiale eve sve ccvovevetereioiclscore srenedelctolews tere MOVeV Ore eter 2a 22 23 27 28 Syl 34 35 36 39 40 47 Figure 16. Linear regression of the abundance and number of species of motile epifauna at each south jetty station as a function oie tenckeul @llewereitem soacoocc00dn000gDD DOD DOU OODDD ODDO OOD GNG000000 17. Estimates of overall mean density for the dominant motile macroinvertebrates of both jetties .......ccccccccccccccccccccees 18. Annual changes in the density of dominant motile macro- invertebrates from the north jetty ......ccccccccrccccccccccccece 19. Seasonal and annual changes in the density of dominant motile macroinvertebrates from the south jetty .......ccccccccsccccccere 20. Vertical distribution of the dominant motile macroinvertebrates Om WOE SCEEHES soocccoon 00D DD DD DD DD DOOD OOOH OUODODOOOHOODDOODDDNN 21. Normal cluster dendrogram of north jetty suction data indicating station groups formed using the Bray-Curtis similarity COGTPULCIEME coodocboDK DD DD OOO ODD OODO DODD DO ONDDUODDDDODDDDOOOOONDDE Table 10. LIST OF TABLES Station designation and date of establishment during jetty CONSEEUCETONU eycrerencvoicicledel elefeneherel ciel ele Gdoddp0OD OOO ODDO OOODOD00000000000 Temperature, salinity, dissolved oxygen and water clarity measurements collected during sampling periods at north jetty SiaAcTons i eereereielee Gd0C0DCODO0U00bDOGG0KS O00000 goodeG00 Oo000000000b00D0N Temperature, salinity, dissolved oxygen and water clarity measurements collected during sampling periods at south jetty GHEANESHOINS Godgagcoc000dD DoDD DU DDDOUOUIOOUC Go000Co0S GO00aD00000D00000 OG Listing of the top ten intertidal and subtidal sessile taxa observed on the north jetty rocks by line-transect census ......... Listing of the top ten intertidal and subtidal sessile taxa observed on the north jetty rocks by photographic census .......... Listing of the top ten intertidal and subtidal sessile taxa observed on the south jetty rocks by line-transect census ......... Listing of the top ten intertidal and subtidal sessile taxa observed on the south jetty rocks by photographic census ...... J00¢ Number of individuals and number of species of each major taxon of motile macroinvertebrates from the north and south WEES. cooocouoGoooocD0b0 a0 booddob0 DDD OGOOOOS DACDDG0DD000 600000000 Species of fishes observed on or near the jetty at Murrells inleesdunineweTreldmstudiles mr O)/9— 19 O2mrariercdeieeeletepensitercrsiereionellcneicrereronenene Percent numerical abundance (N), percent volume displacement (V) and index of relative importance (IRI) of food items found Lie SS EOMach'Stmatererersiererenererenene oooDdOO Dd DODOOCOODOGDONN GoooodDDD000C 17) 18 25 26 32 33 46 58 5) I. INTRODUCTION The South Carolina coast consists of numerous barrier islands separated by estuaries and high salinity inlets. Beach and nearshore sediments in this region are largely composed of sand and shell fragments with very little rocky substrata present. Thus, well-developed intertidal communities of epibenthic organisms are sparse and restricted to the few jetties, groins, and other artificial breakwaters in the area. Subtidal epibenthic communities occur more frequently in association with natural hard bottom areas, artificial reefs, wrecks and jetty rocks, but these habitats are still relatively rare in South Carolina and other southeastern states. As a result, there have been few investigations of the benthos on hard sub- strates in this region, and most of those studies have concentrated on hard bottom areas of the continental shelf (for reviews, see Continental Shelf Associates, 1979; Wenner et al., 1983), or on fouling plate assemblages (Woods Hole Oceanographic Institution, 1952; Sutherland, 1974; Sutherland and Karlson, 1977; Karlson, 1978). Only two studies have been published on the fauna of jetties in South Carolina. Stephenson and Stephenson (1952, 1972) discussed the intertidal biota on rock jetties and breakwaters at Charleston based on a 1947 visit, and McCloskey (1970) characterized the community structure of fauna associated with the coral Ocultna on the Charleston jetties. The Murrells Inlet Navigation Project, authorized by Congress in 1971, provided an opportunity to gain a better understanding of hard and soft bottom marine communities in South Carolina waters and to evaluate changes in those communities following jetty construction. A preliminary assessment of the benthic community at Murrells Inlet was conducted in 1975 (Calder et al., 1976). This report presents detailed data obtained from more recent biological investigations conducted at Murrells Inlet before, during, and after jetty placement. Voiume I describes the colonization and community development of algae, macroinvertebrates, and fish on the jetties. Changes in the nearby intertidal and subtidal infaunal communities are described in Volume II. Specific objectives for the study described in this volume were to: 1. Identify annual changes in the community composition, distribution, and abundance of the algae and macroinvertebrates colonizing the north jetty during the first four years. 2. Document early recruitment and seasonal changes in community composition, distribution, and abundance of algae and macro- invertebrates on the south jetty during the first year, and describe subsequent annual variation. 3. Delineate patterns of vertical biological zonation on both jetties from the jetty base to the supratidal zone. 4. Define differences in community structure related to wave exposure. 5. Identify fish species utilizing the jetty as a habitat, and characterize the food habits of selected species through analysis of their stomach contents. II. DESCRIPTION OF THE STUDY AREA Murrells Inlet, located on the northeastern coast of South Carolina (Fig. 1), is a comparatively small coastal system characterized by ocean beaches, sand and mud flats, intertidal shellfish beds, and expanses of saltmarshes intersected by shallow tidal creeks. Salinities are generally high and stable because of the lack of either a river system flowing into the inlet or contact with the Atlantic Intracoastal Waterway. Water temperatures are more variable, being dependent on the season, and tides are semidiurnal with a mean tidal range of 1.4 m (National Ocean Survey, 1981). At its entrance, Murrells Inlet is flanked by Garden City Beach to the northeast and Huntington Beach to the southwest (Fig. 1). The sediments of these beaches and adjacent nearshore areas consist pri- marily of medium to fine quartz sand with varying amounts of sand-size shell fragments (see Volume II). Although exposed to the open ocean, wave energy is moderate on these beaches because waters are shallow for a considerable distance offshore. Because Murrells Inlet is intensively utilized as the home port for a growing number of commercial and recreational fishing boats, there was a need to stabilize the entrance channel to the inlet. In October 1977, construction began on two quarrystone jetties located on the north and south sides of the inlet entrance (Fig. 1). The north jetty, which extends 1020 m into the ocean, was completed by February 1979. The landward portion of this jetty includes a 411-m weir section (Fig. 1) designed to allow sand to bypass the jetty and settle into a dredged deposition basin, instead of moving around the jetty and creating shoals at the entrance channel. Construction on the south jetty, which extends 1011 m seaward, began in February 1979 and was completed by May 1980. This jetty has no weir section and is topped with an asphalt walkway. Approximate heights of the north and south jetties range from 2.5 to 3.5 m above mean low water (MLW) except at the weir, where the height is approximately 0.7 m above MLW. Crest width on both jetties is approximately 6 m, and the sides slope at an angle of 45° (1V:1H). Granite armor stones of the jetties vary between 5.4 x 103 kg and 9.1 x 103 kg, and individual stone faces vary from horizontal to vertical. Much smaller stones of various sizes are present at the base of each jetty to prevent erosion around the armor stones. MURRELLS INLET OCEAN %S S ww 2 xX | ww a 50 Meters Murrells HUNTINGTON "\ BEACH Figure 1. Map showing Murrells Inlet jetties and station locations. III. MATERIALS AND METHODS 1. Station Characteristics and Sampling Levels Sampling was conducted at four stations on each jetty, two located on opposite sides of the jetty near the outer (offshore) end and two located opposite one another near the inner (inshore) bend of the jetty (Fig. 1). This arrangement provided sampling sites on rocks which had been laid down at different seasons of the year, and also allowed comparisons between wave-exposed and sheltered sides of jetty rocks laid down at the same time. Table 1 provides a listing of the station designations and the date of rock placement at those locations. Six intertidal levels were sampled at each station to provide informa- tion on the vertical zonation of biota on the rocks. These levels were located at mean low water (MLW) and 0.5 m, 1.0 m, 1.5 m, 2.0 m, and 2.5m above MLW. With a mean tidal range of 1.4 m at Murrells Inlet (National Ocean Survey, 1981), these levels encompassed the entire intertidal zone and extended into the supratidal region near the crest of the jetty. Biotic zonation was less pronounced in the subtidal region, and stations were located at 1-m intervals below MLW. Shallow waters in the vicinity of the north jetty limited sampling to levels at depths of -1m (below MLW) on the inner transects and at -1 m and -2 m on the outer transects. Water was even shallower around the south jetty. No subtidal levels were sampled on the exposed side, and only the -l-m level could be sampled on the sheltered side of that jetty. All subtidal levels of both jetties were located at least 0.5 m above the bottom to avoid scouring effects due to wave-entrained sand. By the summer of 1981, additional shoaling had occurred around the south jetty, resulting in burial of the MLW and -1-m levels on the channel side (SPI) and the 0.5-m and MLW levels on the exposed side (SEI). Shoaling continued and by the summer of 1982, the 0.5-m sampling level at SPI was also buried. Benchmarks at known elevations above MLW were marked with paint at the top of each transect. Intertidal levels were located using metered plumb lines oriented to the benchmarks. Subtidal levels were sampled using scuba and were located using a float and metered line. The float line was adjusted to the appropriate length based on tidal height of the water compared to the benchmark height. 2. Sampling Dates The first samples were collected at north jetty stations during July 1979, one year after construction at the inner stations and 8 months after construction at the outer stations. Sampling was then repeated during the summer (July or August) at yearly intervals through 1982. The first south jetty samples were collected in May 1980, seven months after rock emplacement at the inner stations and three months after rock emplacement at the outer stations. Sampling was repeated at 10 Table 1. Station designation and date of establishment during jetty construction. (See Figure 1 for additional information.) STATION LOCATION DATE OF ROCK PLACEMENT NORTH JETTY NEIL exposed side, inner segment of jetty July 1978 NPI protected side, inmer segment of jetty July 1978 NEO exposed side, outer segment of jetty November 1978 NPO protected side, outer segment of jetty November 1978 SOUTH JETTY SEI exposed side, inner segment of jetty October 1979 SPI protected side, inner segment of jetty October 1979 SEO exposed side, outer segment of jetty February 1980 SPO protected side, outer segment of jetty February 1980 11 quarterly intervals (August, November, February) during the first year after construction, and then at yearly intervals (July or August) through 1982. 3. Biological Sampling Methods Characterization of epibenthic communities was accomplished using three systematic sampling techniques: (1) line-transect census, (2) photographed-quadrat census, and (3) suction sampling of motile species. Data collected by the three sampling methods provided information on species composition, relative percent cover or abundance, and distri- bution. In addition, general collections and observations of species were made during all sampling periods. a. Line Transects Percent cover of the sessile biota was assessed at each level using a line-transect procedure modified from Loya and Slobodkin (1971), Porter (1972 a,b), and Loya (1972, 1978). For this assessment a clear plastic strip, marked at its edge with 15 points at 2.5-cm intervals, was placed against rock surfaces. All organisms occurring directly under each point were identified and recorded. Because different rock faces often displayed different densities or assemblages of organisms, assessments were made on each of the seaward, landward, outer, inner, and top surfaces of jetty quarrystone. The transect strip was always positioned horizontally on vertical surfaces, and data from the five rock faces were summed to provide an overall estimate of percent cover based on the 75 points at each level. An effort was made to place the plastic strip on the rock faces without reference to the attached biota to avoid sampling bias. If more than one species was present under a point, all were recorded and percent cover estimates for each species at a given level were based on the percentage of points it occupied. Because this procedure commonly resulted in estimates of total biota cover greater than 100%, total estimated biota cover was determined by substracting the estimated percent of unoccupied space from 100. Poor water visibility and waves precluded in situ assessment by line transect at the subtidal south jetty levels (only). Instead, rocks were removed from the appropriate depth at those stations and brought to the surface for examination. At all stations, organisms which could not be identified in the field were preserved and returned to the laboratory for identification. Samples of blue-green algae were also collected for laboratory identification, but species in this taxonomic group could not be identified in the field and all were identified only as Cyanophyta. b. Photographed Quadrats A photographic census was also conducted to obtain additional quantitative estimates of the jetty epibiota, and to provide a more permanent record of biota at each level. Color photographs were obtained of the same rock faces (i.e., seaward, landward, outer, inner, top) at all station levels using a Nikonos III camera with flash attachment. 2 The camera was equipped with a 35-mm f2.5 Nikkor lens combined with a 35-mm closeup lens outfit and a rectangular quadrat frame (13 x 18.5 cm). As noted for the line-transect census, all faces and levels were located without reference to the attached biota. Photographs were analyzed in the laboratory using a slide projector and a screen with 50 computer-generated random points. One of 10 different screens was selected by random number for analysis of slides from each level. Actual rock surface area examined in each slide was 100 cm2 for the north jetty stations and 150 em? for the south jetty stations. Organisms occurring under the 50 points in each photograph were identified, and percent cover estimates for each species, based on the proportion of points occupied, were calculated for each level (i.e., 250 points/level). Photographic analysis differed slightly from line- transect analysis. Blue-green algae were not assessed in photographs since they could not always be detected, even when present. Furthermore, when there was uncertainty about whether biota existed under a point in the photographs (due to shadows, poor picture quality, etc.), that point was discarded and percent cover estimates were based on the number of analyzable points only. c. Suction Samples Motile epifaunal invertebrates were sampled using a modified underwater slurp gun. The levels sampled were +1 m, MLW, -1 m, and -2 m at all stations, except at the inner stations on each jetty, where shallow depths precluded collection at the -2-m level. Three replicate samples were obtained at all levels by placing the opening of the slurp gun (4-cm diameter) flush against a rock face and vigorously pulling the suction rod. Each replicate consisted of five suctions pooled from different rock faces picked haphazardly. The gun was modified so that suction was obtained by venturi action; incoming water through holes drilled in the barrel was filtered through a l-mm mesh screen. Contents of the slurp gun were emptied into a gallon jug after each suction. To prevent loss of organisms, the mouth of the jug was covered by a 1-mm mesh screen having an opening just large enough to permit insertion of the slurp gun barrel, and the jug was capped except when collections were being added. After the five collections comprising each replicate had been placed in the jug, the container contents were sieved through a 1-mm mesh screen and preserved in a 10% formalin seawater solution. Due to some water leakage around the mouth of the gun and rock face during the suction stroke, the exact surface area sampled per replicate was not defined but approximated 65 cm2. d. Fish Observations and Collections Qualitative observations on ichthyofauna were made during investigations of benthic flora and fauna on the jetty. Fish species observed near the jetties by scuba divers were recorded, baited blackfish traps were set at various locations on the jetty, and a beach seine was pulled along the western side of the weir. In addition, fish species were recorded from gill net collections made in conjunction with a related 13 investigation (Hales and Calder, 1979). Stomachs were removed from the demersal species and preserved for laboratory analysis. In the laboratory, the stomachs were washed in tap water and transferred to 50% isopropanol, and contents of individual stomachs were sorted by taxa and counted. Colonial forms and fragments of animals were counted as one organism unless abundance could be estimated by counting pairs of eyes (crustaceans) , otoliths (fishes), or other parts. Any food items (i.e., fish remains) that might have been bait in blackfish traps were not included in the analysis. Volume displacement of food items was measured using a graduated cylinder, or estimated by using a 0.1-cm2 grid (Windell, 1971). 4. Hydrographic Sampling During every sampling period, surface and bottom water samples were collected at all stations except SEI, which could not be reached by boat. Samples were obtained using a Van Dorn bottle and the parameters measured were temperature, salinity, dissolved oxygen, and water clarity. Water temperature was measured from stem thermometers mounted inside the Van Dorn bottles. Salinity was measured using a Beckman Model RS7B induction salinometer, or a YSI Model 33 S-C-T meter. Dissolved oxygen was measured using a YSI Model 51-B Dissolved Oxygen Meter, or by the modified Winkler titration method (Strickland and Parsons, 1972). Water clarity was measured using a Secchi disk. 5. Data Analysis Community structure was evaluated through comparisons of species cover or abundance, diversity indices, and cluster analysis. Where appropriate, abundance estimates obtained from replicate sampling were statistically compared using the non-parametric Mann-Whitney U test. Only the motile macroinvertebrates were counted since most of the sessile fauna and flora observed on the jetties were colonial. Diversity indices used in the analysis of motile macroinvertebrates included Shannon's index (H') and measurements of species richness (SR) and evenness (J') as described by Margalef (1958) and Pielou (1975). The expressions for these indices are as follows: s H' =-7 a logy a i=1 h where s is the number of species and P; is the proportion of the LE species in a collection, SR = (s - 1) log, n where s is the number of species and n is the number of individuals in a collection, and au = H' logs 14 These measures were computed on data from pooled replicates of suction samples at each level since pooling the replicates provided a larger sample size and a more representative estimate of community diversity at a site. Diversity of the sessile biota which generally could not be counted was limited to comparisons of the number of species (s) observed in photographs and along line transects. Cluster analysis was used to determine patterns of similarity among stations. The quantitative measure used in all analyses was the Bray- Curtis coefficient (Boesch, 1977): estat ey lige Ex + xy) L where X44 and X,4 are the number of individuals of the gen species in two collections under comparison. A normal analysis was completed on the site groups using modified data sets and a flexible sorting strategy with a standard 8 value of -0.25. Data sets represented pooled collections from the different levels at a site (station), separated by seasons. Additional modifications to the data sets included log transformation and deletion of taxa which occurred in only one collection, as well as deletion of those taxa of uncertain identity. These deletions were made to simplify the data sets and because "rare" species usually do not have definable distribution patterns, and can confuse interpretation of cluster analysis. Quantification techniques for food habits of fish are biased, depend- ing on the method (Hynes, 1950; Pinkas et al., 1971; Windell, 1971). Therefore, the relative contribution of different food items to the total diet was determined using three methods: percent frequency occurrence (F), percent numerical abundance (N), and percent volume displacement (V). From these, an index of relative importance (IRI) (Pinkas et al., 1971) was calculated for each prey species and higher taxon as follows: IRI = (N+ V) F where N, V and F are the numerical, volumetric, and frequency percentages as defined above. This index has proven useful in evaluating the relative importance of different food items found in fish stomachs (Pinkas et al., 1971; McEachran et al., 1976; Sedberry, 1983) and was used in the present study to describe the food habits of each species. 15 IV. RESULTS AND DISCUSSION 1. Hydrographic Conditions Water sample analysis for temperature, salinity and dissolved oxygen (Tables 2 and 3) reflected expected hydrographic patterns for this area. Temperature differences between surface and bottom waters were always similar with a normal difference of less than 0.3°C. Lowest temperatures (5.8° - 6.0°C) were observed during the winter and highest temperatures (26.5° - 30.3°C) occurred during summer. Salinity measurements were always high (34.5 - 36.1 °/oo) during the four-year study period since Murrells Inlet receives no significant fresh water input. No salinity data are presented for 1982 due to a faulty meter, but refractometer estimates indicated that salinities were in the same range that year. Dissolved oxygen values were generally high and near saturation values since the shallow waters in this area are well mixed by wave action. Finally, no consistent differences were noted between stations on the north versus south jetty. Water clarity varied considerably during the study, being mostly dependent on tidal stage and wave action. Clarity increased during flood tides and was often greatest on the exposed side of the north jetty. The turbid waters from the inlet decreased water clarity at channel (protected) stations on both jetties, especially during ebb tides. The very shallow waters on the exposed side of the south jetty were also generally more turbid than on the deeper exposed side of the north jetty. 2. Jetty Community Development Data obtained from north and south jetty sampling indicate that a diverse assemblage of biota colonized the rocks during the first four years after construction. At least 25 species of algae, 195 species of macroinvertebrates and 34 species of fish were observed or collected on the jetties, with distinct temporal changes noted each year in the community composition. Vertical gradients in the distribution of fauna and flora on the rocks were also evident, particularly in the intertidal zone. The following sections provide details’on the colonization, community development, and distribution patterns observed on both jetties. a. Sessile Biota. Percent cover estimates for the sessile macroinvertebrates and algal species are listed in Appendices A and B for the four north jetty study sites, and Appendices C and D for the four south jetty sites. Appendix E provides estimates of total biota cover on the rocks using the two census techniques. The line-transect census (Appendices A and C) generally provided more detailed information on community composition at the different levels because taxonomic identifications were often more refined than possible in the analysis of photographed quadrats (Appendices B and D). However, the latter technique did provide useful supplemental information, particularly for the larger dominant biota which could be easily identified. 16 Table 2. Temperature, salinity, dissolved oxygen and water clarity measurements collected during sampling periods at north jetty stations. NEI NEO NPI NPO Surface Bottom Surface Bottom Surface Bottom Surface Bottom TEMPERATURE (°C) 28.4 28.3 28.4 28.4 28.4 28.3 28.3 28.3 28.4 28.2 28.3 28.2 28.9 28.8 28.8 28.8 26.7 26.7 26.5 26.6 26.6 26.7 26.6 26.6 28.2 28.0 28.2 28.2 28.2 28.2 28.2 28.2 35.6 S)3)55) 35.5 ENS}o5) 35.5 35.5 35.5 35.5 S571) 3555 3)3}q5) 35.5 35.4 35.4 35.4 34.5 S56 7/ 35.7 35.6 35.6 35.8 35.8 35.8 35.8 NO DATA DISSOLVED OXYGEN (mg/2) 6.9 7.0 7.0 6.9 6.4 6.7 6.9 6.8 6.7 6.2 6.1 6.0 6.7 6.6 7.0 6.8 6.8 6.4 6.9 6.7 6.5 6.6 6.7 6.8 5.0 5.0 5.1 4.8 4.8 4.6 4.7 4.8 1.9 os} 1.5 1.8 0.7 0.8 1.0 1.0 2.5 2.4 1.6 1.6 17 Table 3. Temperature, salinity, dissolved oxygen and water clarity measurements collected during sampling periods at south jetty stations. SEL SEO SPI SPO Surface Bottom Surface Bottom Surface Bottom Surface Bottom TEMPERATURE (°C) Spring, 1980 24.7 24.1 24.7 24.0 24.2 24.1 Summer, 1980 6 29.3 29/53) 30.2 30.1 30.2 30.3 Fall, 1980 15.7 15.7 15.0 14.9 15}372 15/2 Winter, 1981 i 6.0 6.0 9) 5.9 5.8 5.8 Summer, 1981 A 26.9 27.0 27.4 231/58) 26.9 26.9 Summer, 1982 28.2 28.2 28.5 29.0 28.5 28.2 Spring, 1980 34.5 34.6 34.6 34.5 34.5 34.5 Summer, 1980 5 S5ie 55) S52 Sho? SB}o 72 35.2 Fall, 1980 35a Sie 35.4 35.4 35/5 35/55) Winter, 1981 rl 36.1 36.1 36.1 36.1 36.1 36.1 Summer, 1981 A 35.8 35.8 35.8 35.8 S15) 57/ Sh)57/ Summer, 1982 NO DATA DISSOLVED OXYGEN (mg/2£) Spring, 1980 Use) 7.9 755 7.6 8.0 7.9 Summer, 1980 5 6.9 7.0 Uoal ie) 6.9 6.8 Fall, 1980 7.8 Us?) 8.2 8.3 8.2 8.3 Winter, 1981 i 10.2 10.4 ORE 10.3 10.4 10.1 Summer, 1981 ‘A 6.8 6.4 6.6 6.4 6.9 6.9 Summer, 1982 D2, So? Jo 202 5.4 5.4 WATER CLARITY (m) Spring, 1980 1.4 ale al ib66) Summer, 1980 é 0.9 1.0 1D) Fall, 1980 D 1.0 Io al 1.0 Winter, 1981 : aig al 163} 1.5 Summer, 1981 : 1.6 1.0 Ibe 7 Summer, 1982 NOs DPAU TA (1). Total Biota Cover and Number of Taxa Estimates of total biota cover at the different levels of north jetty stations indicated no consistent or marked differences between inner and outer sites on the same side of the jetty (Appendix E.1). Biota cover on the rocks one year after construction was generally as great as in subsequent years (Fig. 2). This was primarily due to early settling of blue-green algae and the barnacle Chthamalus fragilis intertidally, and settling of the mussel Brachidontes exustus at lower intertidal and subtidal levels. Biota cover at inner and outer south jetty stations did differ considerably in the spring of 1980, with outer sites having less cover than inner sites at all levels where biota was present (Appendix E.2). Rocks at the outer stations had only been submerged for 2-3 months as compared to 7 months of submersion at the inner stations. By the summer of 1980, biota cover at all levels on the rocks had increased to percen- tages as great or greater than those found in subsequent sampling periods (Fig. 3). In the upper intertidal zone (1.5 m - 2.0 m above MLW), biota cover was often greater on the wave-exposed side as compared with the sheltered side of the jetties (Figs. 2 and 3). This vertical extension in the amount of biota cover on the exposed side is a common pattern which has been observed in several other rocky intertidal systems (Lewis, 1972). Biota cover on the rocks of both jetties generally increased at the lower levels, and differences between sides were not as great. Because cover on the exposed side was rarely less than on the wave- protected side (Figs. 2 and 3), it is unlikely that wave shock represents a major source of mortality as noted in other rocky intertidal systems (Dayton, 1971; Menge, 1978). However, wave energy in those systems is often considerably greater than the moderate wave energy observed at Murrells Inlet. Comparisons of biota cover estimates obtained by line-transect versus photographic census (Figs. 2 and 3) showed strong similarities except at the highest intertidal levels. Blue-green algae were dominant in the upper inter- tidal zone, and these species of algae were not assessed in the photographs. Since many of the sessile organisms are colonial, species diversity indices were not calculated on this component of the jetty communities. However, an examination of the number of taxa found on the rocks indicates that there were fewer species in the intertidal zone than in the subtidal zone on both jetties (Figs. 4 and 5, Appendices A-D). The more rigorous physical environment associated with the intertidal habitat obviously limits the number of species which can colonize this area as compared with the less stressful subtidal environment. In both the intertidal and subtidal zones, the number of taxa present on the jetty within one year after construction was nearly equivalent to or greater than the number found in later years (Figs. 4 and 5). Additionally, there were no major or consistent differences in the number of taxa found on the wave-exposed versus sheltered sides of the jetties. 19 ‘elep ou SOJPOTPUT yxy Spoaquesoidoa’ st uoT}vjs ouo ATUO Jey SozeOTpPUT » ‘apts (d) peqz0e301d 9a pue apts (4) posodxd oy} uO suOT}eIS OQ YQ WOT; soSeisae quosoidez sweiasoqsty °AqQof YyY_Aou oYyR JO STaAZT JUaeADFIZIP qe AJOAOD BIOTA STTSSes Jo sojzeuTIso yderasoqoyd pue JoOssueAR-OUT]T °*7 VANSTY 2861 186l os6l 6261 286i IS6l Os6l 6261 Jsawwng sewwns sowwe: seWWNS Joewwng sewwns Joawwng sewwns d 3 d 3 d 3 d 3 0'2- [|| [|| [|!- MM 4 OM % OH % Or * % OH % * Loot O'- os os oot oo & 00 mi D os os v3 m 001 oo! % % s'0 m D OS o O m 2 001 oo Ol @ | | | | ill il os os O s oo! 001 G : ° Ie 2 "WIL. 1Jbos os 5 001 001 os os 001 001 ae sz Se Se ee oa At) os TV3A31 SHdVY9SOLOHd SLOASNVYL SANIT 20 SoqWeoTpUuT xx ay. JO STXAST JUIAIFJTP Je A9AOD BROT eT}TSses jo 2861 1861 os6! 086) O86! O86) JOWWAS JOWWNS JOIUIAA Wo4 JOwWWNS 6uysds d 3 d 3 d 3 d 3 d 3 d 3 - RK iB i: x its OF * ¥ aR * % Me -OS M. 00! ee si % ve < M as % ™ oo! | | cers GO yy wo, Bu il] | | yy 2 ol OS x ool BwW W =i. ru St OS = = 00) a a % seat 0S = 001 eee er ras | 2 (w) OS 1V3A31 SHdVYU9OLOHd ‘pajueseidez st uot}eqs suo ATUO Jeu Se eDTpUT yx apts (q) pesodxe oy} uo suot}ze}S OM] DY} WOTZ Sese1eAe YUeSeide1 swe130ISTH ‘eqep ou ‘ePTS (d) paqoeqoid ay. pue *£3Q9ef y Anos sajeuytise ydersoj0yd pue Joosuezq-9UTT ‘+e 9ain3Ty 286! 166) 1861 os6l O86! O86! JOWWNS JOWWNS JOUUIMA WO4 JQWWNS bulsds d3 d 3 d 3 d 3 d 3 d 3 ¥ ¥ ¥ ¥ aie} %O% * * * * % I l I Wye 4. THOU Ye -Oo1 ay Wy wy. so ay a i Y3AOD VLOIG LN39¥3d a0VeaAV SLOSSNVYUL SANIT 21 Intertidal e———e Exposed Side o- —- — © Protected Side Subtidal NUMBER OF TAXA JAN. JAN. JAN. JAN. I978 i979 1980 I98i 1982 Figure 4. Total number of sessile taxa observed at intertidal and subtidal levels of north jetty stations during line- transect census. 22 Intertidal ¢ x< 2 o——— Exposed Side Le Oo —— © Protected Side Oo or Subtidal J a = = =z JAN. JAN. JAN. 1980 I98\ i982 Figure 5. Total number of sessile taxa observed at intertidal and subtidal levels of south jetty stations during line-transect census. 23 If wave stress at Murrells Inlet had been greater, differences in the diversity of species might have been more apparent, as noted in other rocky intertidal systems (Menge and Sutherland, 1976). (2). Community Composition (a). North Jetty Although estimates of total biota cover and the number of taxa on the north jetty rocks did not change markedly over the four-year study period, community composition of the sessile biota did vary considerably between years (Tables 4 and 5; Figs. 6 and 7). Major differences were also observed in the intertidal versus subtidal community composition and, to a lesser extent, between the wave-exposed and protected sides. One year after jetty construction, the dominant intertidal species at all stations were the barnacle Chthamalus fragilis and the mussel Brachidontes exustus. These two species accounted for approximately 63% of all biota cover in this zone. The oyster Crassostrea virginica and the barnacle Balanus eburneus were the only other fauna among the ten dominant organisms found intertidally. Blue-green algae (Cyanophyta) was the primary intertidal algal form during the first year. The dominant species of blue-greens were Anacystts aeruginosa, Microcoleus lyngbyaceous, and Calothrix crustacea. These three species were noted during all later sampling periods, as well. Other algal species found on the rocks included the green algae Cladophora sp. (primarily C. laetevtrens) and Ulva sp., and the red algae Hypnea musctformis, Lomentarita batleyana and Herpostphonta tenella. Although barnacle and mussel cover was generally similar on both sides of the jetty in 1979, blue-green and green algae were the predominant algal forms on the exposed side, whereas red algae were predominant on the protected side. Chthamalus fragilis, Brachtdontes exustus, Ulva sp. and Cyanophyta continued to dominate the intertidal biota cover during the next three years (Tables 4 and 5). In 1980, rock coverage by the different taxa was similar to that observed in 1979 (Figs. 6 and 7), but by 1981, algal cover had increased. The line-transect census indicated that blue-green algae was more prevalent this year than in any other year. Larger macrophyte coverage had also increased to a lesser extent, with green algae (Ulva sp., Enteromorpha sp., and Cladophora sp.) generally being more common on both sides of the jetty than red algae (Gracilaria folitfera, Porphyra sp., Hypnea musctformis and Polystphonta sp.). By 1982, algal and mussel cover had decreased considerably. Barnacles (C. fragilis) represented the dominant biota on the rocks, although blue- green algae was also common. The decline in algal and mussel cover during the last year of study is not readily explained. Some of this decline may be attributed to mortality. Additionally, both taxonomic groups were concentrated in only the lowest portion of the intertidal zone during all four years, and the MLW sampling level represented the upper limit for many of the 24 ‘ds paoydopv7) pa1y4,0b 011 et0doum1 DY, ‘ds pyaoydopv1) sisuazqpyunu D7nNB2Z oY OT supjsip D1IUDINVWeg Du141deu vynbng pupha] 1g DLIDZUaUOT purorutos Dueumuhiq 6 sisuepnudeg D141 db}87q ‘ds p1s1u9 piaf1110f DLb]10vAN 1iseqaiof sp1da7qsy 8 pyourbuou D1UDINZAAS ‘ds n2ucydishjog asueu1jzodpo DUOISTpNy ‘ds umip17 dy L psoun}d s1isdohug snysnxe saqyUuopiyobug pu14ziieu vynbng Duvhe1 1g DIWDZUSUOT 9 pyojno1ueBb v2172q0 sisuapniuteg 017 d0781q snqsnxa saqzuop1yobug ‘ds v2uoydrshz0g S asuaUurjoxpe DUWOZSTpPNY puozoya1p 02172q0 DyvdIda D1] atodoz1yag pur4idiau pynbng 0 sqmiofrosnu vaudhy pyzouyodopnasd pviuauhpoyy ‘ds Da7n sip2a1a v1oydoredg € pyouzodopnasd prueuhpoyy ceeerOn DLID1 1LODAD pvypuzodopnasd piruauhpoyy puozOYyoLp 011 2q0 G pdaf1110f D1aID] 1DID aSUuaUu1]Oubo DUWOZSIPNY puozoyo1p D0272qG0 snqsnxe seqzuop1yopug T TVOILANS snsi¢crdur snup] Dg pypuypodopnasd piuauhpoyy siulofiosnu poudhy “ds va7n ot snq:1uaa snup1 Dg ‘ds piuoydishj10g sns1aoudut snub] Dg ‘ds vaoydopv79 6 vsounid sisdohug *ds vahyditog ‘ds vuaoydopr79 D1] 0Uey DiLUoYydisodizey 8 snauinga snun] pg po1urbita vedzsossvdag ‘ds py diowo1ezUg puvhia]1pq D1WdAUAUOT l simiofiosnu Deudhy puaf1110f D1uDp] 10bA)N pO1Utbi11 DatzsossvsA) snauinga snuv] pg 9 DOLULIBALA DadtZSOSSDAQ ‘ds pnyditowo1equq ‘ds vauoydishjz0q DOLUIBALA DadtZSOssDdg S snisnxa saquop1yovoug ‘ds va7zn ‘ds Da7N squiofiasnu veudhy ) ‘ds Da7n 8111bbaf snzpumyzy) eihydoueky e 3 Aydouedy € e3 Aydoued9 snysnxe seqzuop joDug snqsnxe saqzuop1lyopug 8217 1boaf sn] DumDYy2Yy) z $21 21bbaf snj7oumyzy9 eq) .ourt) s271bpaf sn. pumyzYyo snqysnxe sazuop1yoDpdg T TWAT LYAINT 786L T86T O86T 6261 INVY YaWWNS YaWWNS YaWWNs YaWWNS *snsu9. 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Figure 6. Histograms represent means from the 2.0-m to MLW intertidal levels, and the -l- and —2-m subtidal levels. 27 Exposed Protected INTERTIOAL Ce 1981 60 1981 40 20 20 g AVERAGE PERCENT COVER Barnacies Mussels Tigure 7. $ 60 1982 1982 20 SUBTIDAL 1979 20: 1962 K>»éSGW] 23 | eeeoes9se## ese e®ee#eed$dtdgesd #8.» tw e¢ @ oo= Ft 98 c¢ @® 533225 $3538 285 2 OS ES se e@ad¢eEFsa2o0 sss ae FE B35 Oe Se = 3 0 ZVes s‘uvu2e@e e a> @a -¢ = < a ez < co) o = Photographic estimates of mean percent cover for tne different sessile taxa found on the north jetty rocks. Wistograms represent means from the 2.0-m to MLW intertidal levels, and the -l- and -2-m subtidal levels. species. During the 1982 assessment, algae and mussels were abundant just below the MLW sampling level. Thus, it is possible that the levels selected in that year were slightly higher (2-5 cm) than the levels selected in previous years, causing lowered estimates of these taxa. In the subtidal zone, different taxa dominated when compared with the intertidal zone, and major changes occurred yearly in the most abundant biota (Tables 4 and 5, Figs. 6 and 7). One year after construction, the subtidal rocks were primarily covered with the mussel B. exustus on the exposed side. Rocks on the wave-sheltered side had fewer B. exustus and more ascidians (primarily Perophora viridis, Aplidium sp. and Molgula manhattensts), but mussels were still the dominant taxon. One year later, mussel density had declined significantly and both sides had heavy algal cover. The decreased mussel density was most likely due to predation by sheepshead (Archosargus probatocephalus) and by dense aggregations of the starfish Astertas forbestt, which were seen grazing on the mussels during the summer of 1979. Mussel density remained low on the subtidal rocks for the remainder of the study. Prominent algal species which colonized the rocks during the second year included the red algae Gracilaria folitfera and Rhodymenta pseudopal- mata, and the green alga Ulva sp. Algae appeared to be more prevalent on the exposed side, which may be due to greater light penetration in the clearer waters generally observed on that side. Bryozoans were also common on the exposed side in 1980, and they were much more abundant there than on the wave-protected side. Dominant bryozoans included the stalked forms Angutnella palmata and Bugula nerttina, and the encrusting form Schtzoporella errata (Tables 4 and 5). On the protected side, hydroids and ascidians were more prevalent than bryozoans, which were relatively rare on that side. The most common hydroid was Obelta dichotoma. This species grew primarily on the red algae. The dominant ascidian was Eudtstoma carolinense, which formed large mats of clumped zooids having sand-covered tests. Because of its morphological complexity, this ascidian species provided excellent habitat for a diversity of smaller, more motile invertebrates. By 1981, the third year after jetty construction, ascidians had become quite common on both sides of the jetty. Eudtstoma carolinense and Distaplia bermudensis were the most abundant ascidians on the exposed side, and E. carolinense remained as the dominant species on the wave-sheltered side (Appendices A and B). Algae was also common on both sides, with red algae (G. folttfera and Rk. pseudopalmata) being more prevalent than green algae. The hydroid Obelta dichotoma was often observed growing on the red algae. Many stalked bryozoans, such as Crista sp., Angutnella palmata, and Bugula neritina were commonly observed attached to the rocks. The encrusting bryozoan Sehtzoporella errata did not appear to be as common in 1981 as in the preceding year, but another encrusting forn, Thalamoporeltla gothica, was noted for the first time on the rocks (Appendix A). This latter species often grew in large erect colonies shaped in the form of lettuce heads. Finally, the octocoral Leptogorgia virgulata, was often noted growing on the rocks, especially at the base of the jetties. 29 During the last year of study, dominance in biota cover had changed once again. Although ascidians were still common, especially on the channel side of the jetty, algae represented the dominant biota cover in 1982 (Tables 4 and 5, Figs. 6 and 7). Red algae was much more prevalent than green algae, with lush stands of Rk. pseudopalmata, G. folitfera and H. musctformis covering the hard substrata. Rhodymenta pseudopalmata was more abundant on the exposed side, and it was the most prominent alga in the shallower depths (-1 m). Hydroids, such as Dynamena spp., Obelta spp., and Sertularta spp., were common on the algae and rocks of the exposed side. Hydroids were also common on the channel side of the jetty, but to a lesser extent. However, one hydroid (Halocordyle ditsttcha) appeared to be more abundant on the channel side than on the exposed side. Eudtstoma caroltnense and D. bermudensts were the most common ascidians on the rocks during the summer of 1982. Cluster analysis of line-transect data on the entire north jetty sessile community (Fig. 8) supports the hypothesis that the overall community structure changed substantially over the four-year period. All stations sampled in the first year grouped together with a relatively high degree of similarity in faunal and floral composition (Group 1). Within that group, the inner stations were more similar to each other than to the outer stations, indicating differences in community composition between those station groups. These differences were probably related to the 6-month difference in time of rock submer- sion and the difference in the number of levels sampled at the outer stations. With one exception (NPO; SU, 1980), community structure in 1979 was relatively dissimilar to the community structure observed in later years (Groups 2 and 3). Station collections from 1980 and 1981 formed Group 2, demonstrating more similarity among collections from those years than from the first and last years of the study. Within that group, 1981 collections were generally more similar to each other than to 1980 collections. The two 1980 collections at the inner stations remained relatively dissimilar to the other collections in that year and in 1981. Community structure at stations sampled in 1982 was very low in similarity to the same stations sampled in previous years (Group 3). Additionally, there was a distinct separation of protected and exposed stations based on faunal similarity. These major shifts in overall community composition between years, combined with the documented yearly changes in dominant biota cover (Figs. 6 and 7) strongly suggest that the sessile community on the jetty rock had not stabilized by the fourth year after construction. (b), South Jetty Species composition and structure of the sessile community on the south jetty also changed considerably between sampling periods (Tables 6 and 7, Figs. 9-11). Additionally, many of these changes differed from those noted on the north jetty, primarily because this jetty was sampled seasonally during the first year after construction. The initial May 1980 assessment documented that the intertidal rocks were still relatively devoid of fauna, and algal coverage in that zone was only moderate (Figs. 9 and 10). The dominant algae were Porphyra sp. 30 Group Season Station SU 82 NPI SU 82 NPO 3 su 82 NEI SU 82 NEON hes a 3 SUreO? NEL ae SU 80 NPI SU 80 NEO 2 SU 8 NEO SU 8l NEL SU 81 NPI SU 81 NPO SUS yo WEL po a SU 79 NPI | SU 79 NEO SU 79 NPO SU 80 NPO 8 6 4 2 fo) SIMILARITY Figure 8. 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Line-transect estimates of mean percent cover for the different sessile taxa found on the intertidal south jetty rocks. Histograms represent means from the 2.0-m to MLW intertidal levels. 34 AVERAGE PERCENT COVER Exposed Protected SPRING 1980 60 SPRING 1980 40: 20 SUMMER 1980 60 SUMMER 1980 40 20: wm £ OO -’ Oo 5 0 0 8 o 38 60 FALL 1980 6 FALL 1980 40 40 20 20 60 WINTER 1981 60 WINTER 1981 40 40 20 20 IN| UJ 60 SUMMER 198! 60 SUMMER 198! 40 40 20 20 60 SUMMER 1982 60 SUMMER 1982 40: 40: 20 20 a /-he, otee oe oe se (e Seo 25 Sh ob ono. = Se oS SSSR RS E Sie -Sin Suet Seek Sis ee ee ee chisita fcS Sa me 10 o=O° ce vo >D B B'S eS 5 eo oO < P3 = qd o < 8 =x < © o Figure 10. Photographic estimates of mean percent cover for the different sessile taxa found on the intertidal south jetty rocks. Histograms represent means from tne 2.0-m to MLW intertidal levels. 35 AVERAGE PERCENT COVER Line Transects Photographs SPRING 1980 60 SPRING 1980 40 20 Barnacles Mussels Figure 11. Oysters 60 SUMMER 1980 FALL 1980 ee FALL 1980 WINTER 1981 60 WINTER 198! SUMMER 1981 SUMMER 1981 o so e $S8$0e980€e 86 *©35£ss 38 € & EE EAs Ses i SO Sao oe szomo 8B 65 #®= Oc >D B oo oe = q @ eo eo = dq a 5 1 © ° @ 2 @ Line-transect and photographic estimates of the mean percent cover for the different sessile taxa found at the -1.0-m subtidal level on the protected side of the south jetty. 36 and Enteromorpha sp. (Tables 6 and 7). Kapraun and Zechman (1982) also noted these genera colonizing North Carolina jetty rocks during the winter and early spring months. Two barnacle species, Chthamalus fragtlts and Balanus improvisus, the hydroid Tubularta crocea, and the bryozoan Bugula neritina were the only fauna found on the rocks in May. Larval recruitment of these species is known to begin in the cooler water temperatures which prevailed at Murrells Inlet during the period preceding first sampling (Woods Hole Oceanographic Institution, 1952; Sutherland and Karlson, 1977), thus accounting for their presence. Three months later biota coverage had increased considerably. The most abundant three organisms (Cyanophyta, C. fragilis, and B. exustus), which had settled on the rocks during the intervening period, were the same as those noted on the north jetty at that time (Tables 4 and 6). Chthamalus fragitlts and blue-green algae were more prevalent on the exposed side during that summer, but by autumn, this difference between sides was reduced. Mussel coverage also increased during the summer months of 1980, reaching peak densities on both sides by November (Figs. 9 and 10). Mussels, barnacles and blue-green algae continued to dominate the community in terms of rock coverage until the summer of 1982 when coverage by most taxa declined. Reduced biota coverage on the rocks in 1982 reflected, in part, the reduced number of intertidal levels which could be sampled in that year. For example, coverage of the intertidal rocks by the mussel B. exustus appears to be considerably less in 1982 as compared with the preceding sampling periods (Figs. 9 and 10). However, this species was observed only in the lower portion of the intertidal zone on both jetties, and these sampling levels were buried in sand at the inner stations (SPI, SEI) by 1982. Additional causes for the reduced biota cover observed on this jetty may include such factors as competition, predation, and natural mortality. Even at stations where the lower levels were not buried, mussel densities had declined considerably at the lower levels by 1982 (Appendices C and D). Additionally, although the rocks at higher intertidal levels appeared to be covered with barnacles, close inspection revealed that most were just shell plates from dead adults and the majority of living specimens were newly settled juvenile forms, The natural life span of the dominant species, C. fragilis, is not known for this area. However, a 2- to 3-year life span has been noted for other barnacle species (Woods Hole Oceanographic Institution, 1952), which correlates well with the mortality noted on the south jetty. As noted for the north jetty biota, subtidal fauna and flora on the south jetty rocks changed considerably over the study period (Tables 6 and 7, Fig. 11). Three months after completion of rock emplacement, several species had colonized the rocks, but the hydroid Tubularta crocea dominated in terms of faunal cover. The peak settling period for this species is during the spring (Woods Hole Oceanographic Institution, 1952; Sutherland and Karlson, 1977), thus explaining early dominance on the rocks. By summer, 7. crocea had disappeared from the rocks, probably because this species undergoes cycles of activity in temperate areas of the western Atlantic, and it is inactive during the summer in South Carolina (Calder, unpublished). 37 During the summer of 1980, the mussel Brachidontes exustus covered more than 50% of the subtidal rock space at station SPO, whereas the bryozoan Bugula neritina and the red algae Lomentaria batleyana were more prevalent at station SPI (Appendix C). These differences probably reflect the different duration of rock submergence at these sites as noted for the north jetty stations. Other fauna commonly found in the subtidal zone during this season included the polychaetes Sabellarta vulgarts and Hydrotdes sp. (mostly H. dtanthus), the ascidians Molgula manhattensts and Perophora virtdis, the barnacles Balanus spp., and the encrusting bryozoans Schtzoporella errata and Membrantpora tenuts (Tables 6 and 7). Mussels continued to dominate the subtidal rocks at SPO for the remainder of the study, generally covering more than 70% of the rocks. At SPI, on the other hand, ascidians, serpulid polychaetes, and hydroids formed the dominant biota cover (Appendices C and D). With few exceptions, the subtidal community composition at both south jetty stations was not very similar to equivalent areas on the north jetty during the same sampling periods, or after equivalent periods of rock submergence. This was particularly evident for the algal component, which dominanted biota cover on the north jetty rocks during 1981 and 1982, but not on the south jetty where algae were rarely observed. Cluster analysis of south jetty data confirms that some seasonal and yearly changes occurred in the jetty community composition (Fig. 12), but the differences are not as clear as those noted on the north jetty. Stations sampled during the first season (Group 1) had relatively dissimilar faunal and floral composition to all other station collections. As noted previously, biota cover on the rocks at this time was relatively depauperate, thus accounting for this separation of collections, Station groupings from most later collections did not indicate any distinct seasonal separation (Groups 2-4), but it is interesting to note that within those groups, collections from the protected side often grouped separately from collections on the exposed side. This is most probably due to the presence of subtidal fauna and flora at the protected sites as compared with the exposed side where waters were too shallow to sample subtidally. (3). Vertical Zonation Patterns Obvious gradients were observed in the vertical distribution of most species found on the north and south jetties (Figs. 13 and 14). These distribution patterns were generally similar over the entire study period, with only minor differences noted between sides (Appendices A-D). Rocks at 2.5 m above MLW were usually devoid of any biota since this level was well above the mean high water mark. Only occasional small patches of blue-green algae were noted. These patches were probably established during periods of heavy wave swell (e.g. storms), but they appeared to be short lived based on observations during subsequent sampling periods. 38 *quoToOTFZyooo AQTAeTTWES stqin9-Aeig ay} B3utsn poewz0j3 sdnoz3Z uotzeISs But IeOTpUT eqep qoesueaj-outyT Aqjef y3nos jo stsdkTeue 1z93SNTD TeWION “ZT oInsTtyq ALIYV TIWIS = €- = iP u01s01S uospes dnoi9 39 *potaod zeaf-1nojF vy} AVAO suoT}eqS [Te worF senyTea uesu Juesaiderz sajewtqysyY *suotjeqs AqJel yZA0U J® poAresqo sotoeds oTtsses Juepunqe Jsow QZ 24} JO VOTINGTAISTP TeITIAOA “ET VaNBTy y ¢ PE : i g j i 3 e md a 8 ah, 3 ‘ w Hy OS 8 : Ly. y P Te iy & i aie : eee. oe ‘ 8 i phys Gd of y pe org & fa : ae se pele es g a 4 &. S = g 5 gee a Vs CS S 3 a x 3 & e Ry = 3 & 3 Pees 3 A : s & aed & $ § g ee of g & 3 2 g 3 E gsis3 3 tee td es _ sydosbojoug W VV Ae SN ON Ye Ne ree VvOZOAN8 V3OVIGIOSY VOZONGAH VOSNTIOW SJIOVNYVE 3V91V (W) 13A37 40 *potied zeeaf4-901Y} 24} 19A0 suoTqe3S [[e wory senTea ueeu Jueseadea sejeuT3sy <‘suotzeys AqJef YInoOS qe paaiesqo setoeds e[Tsses Juepunge jsow QZ 94} JO UOTINGTAISTP TeOTIAGA HT ain3stq 5 Hy g OR d 2 9 j We oi ee es : ‘ ; BG ip fF ite ale EAS 1S | Lees a! a8 ay 9 2 a zt } E i § Ss Gs cS) 5S Po Rd : 3 " reaee RB bs : fe oy : = i ‘ He GBetp BR Bo 8 H pO OB Rg a 8 : 3 z mgag (Bb g 3 2 ee ee eS eS eee —— EE Paes 60 ~ 1- ie) 1+ Ror = 21V9S sydosboyoug oi $j998UDJ, OUI Ne fg NS NEE a NS VOZOANS) 3=V39VIGIOSV VOZOUNGAH VOSNTION S31DVNUVE av91V (Ww) 13A31 41 Blue-green algae cover increased considerably at the next two lower levels, becoming the dominant biota cover at 2.0 m above MLW (Figs. 13 and 14). As noted previously, the common blue-green algal species observed on the rocks throughout the study were Microcoleus lyngbyaceous, Calothrix crustacea, and Anacystis aeruginosa. The combined cover of these species was usually dense enough to form a thick "black-colored" band on the rocks. The only macroinvertebrates noted at 2.0 m above MLW were the barnacles Chthamalus fragilts, but average coverage of this species was quite low at this level. Chthamalus fragilis density was much greater at 1.5 m above MLW and this species was often the only fauna present on the rocks at this level. The barnacles were usually coated with blue-green algae which often ob- scured them when observed from a distance. Other algae occasionally observed at the 1.5-m level were Porphyra sp. and Enteromorpha sp. Maximum densities of C. fragilts were observed at 1.0 m above MLW. At this level, other species which had been either rare or absent at higher levels were also present on the rocks. These included the mussel Brachidontes exustus, the oyster Crassostrea virgintca, and the algae Ulva sp., Enteromorpha sp., Polystphonta sp., Porphyra sp. and Cladophora sp. Only Porphyra sp. reached its maximum cover at this level. All other species were more common at lower intertidal levels. Although blue-green algae were still present at 1.0-m above MLW, their percent cover was considerably reduced as compared with their coverage at higher levels. Major changes occurred in the sessile community between the 1.0-m and 0.5-m levels, primarily due to a change from barnacle to mussel dominance. During the earlier stages of jetty community development, the 0.5-m rocks were covered by dense mats of B. exustus and numbers of C. fragilis were greatly reduced. Although mussel density decreased during later years, mussels were still abundant at this level and at MLW. Chthamalus fragilts was rare or absent at the 0.5-m level during the first sampling period on both jetties, but more numerous later on the bare rock space that was opened when mussel density declined. Another barnacle, Balanus eburneus, was also present on the rocks at 0.5 m and the oyster C. virginica was most common at this level (Figs. 13 and 14). The number of sessile species attached to the rocks increased substantially at the MLW level (Appendices A-D), although mussels still heavily dominated the rocks. When mussel density declined with time, algal growth on the rocks increased and generally filled the bare rock spaces. Species which reached peak abundance at this level included the green algae Ulva sp. and Enteromorpha sp., and the red algae Hypnea musctformis, Polysiphonta sp., and Gracilaria folitfera. Blue-green algae were least abundant at this level and were rarely observed below MLW. Oysters and barnacles were also common at MLW but C. fragilis was replaced by Balanus spp., primarily B. eburneus. Gradients in the vertical distribution of most species found sub- tidally were less pronounced, mostly because there were fewer differences in the physical environment between levels of that zone. However, it is 42 important to note that many of the taxa observed at subtidal levels, such as hydroids, bryozoans and ascidians, were present in only that zone. This is due, in part, to their inability to tolerate desiccation and other stresses present in the intertidal zone. In the first year, B. exustus was extremely abundant on the subtidal rocks, particularly at the outer stations which had been submerged for the least period of time. As noted previously, when mussel density declined, algae, ascidians, bryozoans and hydroids became more prominent on the rocks at both levels. Generally, green algae were more prevalent at the shallower depths where better light penetration occurred (Fig. 13). Red algae species were also common at that level and at the -2-m level where green algae were rarely observed. Most hydroid, bryozoan, and ascidian species were also slightly more common at the -2-m level where algae cover on the rocks was not as dense. Some exceptions to this trend were the hydroids Obelta gentculata and 0. dichotoma, which were most often observed growing on the red algae G. folttfera and R. pseudopalmata. On the channel side of the rocks, light penetration was generally lower and algae were not as abundant. As a result, hydroids, bryozoans, and ascidians were more common at shallower depths on that side than the wave-exposed side (Appendices A-D). (4). General Discussion Community composition and patterns of vertical zonation resembled those described from the jetties at Charleston, South Carolina, by Stephenson and Stephenson (1952, 1972). They reported finding the barnacle Chthamalus fragilts in the black band of blue-green algae and lichens marking the splash zone (supralittoral fringe). Peak abundances of this barnacle were found high in the intertidal zone. Mussels (Brachidontes exustus), reported in "colossal quantities," oysters (Crassostrea virgintea), and balanid barnacles (Balanus eburneus, B. improvisus) were the dominant invertebrates in the middle and lower intertidal zones. Although mussels and balanid barnacles were also abundant near MLW on the Murrells Inlet jetties, oysters were not always common. Other invertebrates reported near MLW on the Charleston jetties included the gastropod Urosalpinx cinerea, the scleractinian coral Oculina arbuscula, the ascidian Molgula manhattensis, the asteroid Asterias forbesti, the echinoid Arbacia punctulata, the hydroid Tubularia crocea, the bryozoans Angutnella palmata, Electra monostachys, and Membrantpora tenuis, the actiniarian Bunodosoma cavernata, and a red sponge believed to be Hymentactdon heltophtla. Many factors, both biotic and abiotic, influence epifaunal community structure and development on rocky shores. Although biotic factors have not been ignored, most descriptive investigations have attributed distributional patterns largely to abiotic factors such as wave exposure, tides, desiccation, climate, temperature, light intensity, width of the rocky tract, proximity of sand, and rock composition, texture, and configuration (see Stephenson and Stephenson, 1972; Lewis, 1972; Newell, 1979). Experimental research, beginning with Connell (1961a,b; 1970) and including studies such as those of Paine (1966, 1969, 1974), Dayton (1971, 1975), Menge (1976), Lubchenco and Menge (1978), and others, has emphasized biological interactions such as predation and inter- and intra-specific competition in community development and species distribution. 43 Such studies generally support the hypothesis that the lower limits of intertidal species are mainly biologically controlled while upper limits are more likely set by abiotic factors (Connell, 1972). Lewis (1977) cautioned that neither biological nor physical factors should be under- estimated in the distribution of rocky shore communities. At Murrells Inlet, physical stress most likely controlled the upper limits of Chthamalus fragilis and blue-green algae; no evidence of significant barnacle predation was observed at the higher levels. Although not tested, we believe the lower distribution of C. fragilis may be limited by competition for space with B. exustus, which formed dense mats of biota at 0.5 m above MLW. The relative distribution of both these species parallels that noted by Menge (1976) for Balanus balanotdes and Mytilus edults in a New England rocky intertidal system. Even when mussel density declined with time, other dominant forms replaced mussels on the bare rock space and it is likely that C. fragilis was still competitively excluded. The upper distribution of the mussels B. exustus and algae Ulva sp., Hypnea musctformis and Gracilaria foltifera also appeared to be regulated by physical factors since the mat of mussels stopped abruptly just above the 0.5-m level and the algae were rarely found above MLW even though there was space available for all these species to colonize the rocks. Within the intertidal and subtidal zones covered by B. exustus, mussel density may have been influenced by biotic factors. Intertidally, large numbers of birds, particularly overwintering ruddy turnstones and various species of gulls, were observed on the jetties during February. These birds were seen feeding around jetty rocks, and shell fragments of B. exustus were abundant in bird excrement on the jetties. Subtidally, the mussel predator Astertas forbesti was often observed on the rocks, apparently feeding on B. exustus. Therefore, it is likely that the decline in mussel density after the first year was due to predation since the natural life span of most mussels is longer than one year (Woods Hole Oceanographic Institution, 1952) and mussels are generally competitive dominants in rocky intertidal systems (Paine, 1974; Menge, 1976). In terms of the overall sessile community composition on the jetty rocks, species composition and vertical distribution patterns in the more physically stressed intertidal zone appear to have approached relative stability quickly. The well-defined bands of blue-green algae, barnacles, oysters and mussels were established within the first 12 months after rock emplacement, and alterations in invertebrate community structure were relatively minor thereafter. Subtidally, epibenthic communities appeared to be less stable over the four-year study. Mussels, which initially dominated the subtidal community, were replaced by bryozoans, ascidians, cnidarians and algae with major changes occurring in the yearly dominance of taxa. Additionally, differences in community composition were observed between jetties in subtidal areas sampled during the same season, and even between sides on the same jetty. These differences were probably due to differences in time of rock submersion and wave exposure (Woods Hole Oceanographic Institution, 1952; Calder and Brehmer, 1967; Connell, 1972; Osman, 1977). However, the observations during this study support Sutherland's (1974) and Sutherland and Karlson's 44 (1977) contention that a stable "climax" community of sessile invertebrates is not likely to occur. b. Motile Epifauna Ranked abundance estimates for all motile macroinvertebrates are provided in Appendix F for the four north-jetty sites and in Appendix G for the four south-jetty sites. (1). Total Abundance and Number of Taxa Slurp gun sampling at intertidal and subtidal levels of both jetties resulted in the collection of 131 species over the four-year period. Amphipods, polychaetes and molluscs represented over 70% of all species found on the north-jetty rocks (Table 8). Amphipods alone dominated the motile epifauna on the south jetty, representing approximately 40% of all such species collected. On both jetties, motile biota rapidly colonized the rocks, with densities generally as high in the first sampling period as in subsequent sampling periods (Appendix H). Amphipods and isopods were the two most numerically abundant taxa on both jetties even though only a few isopod species were represented. Over all levels, 11 major taxonomic groups were found on the north jetty and 10 on the south jetty (Table 8). Both the number of species and the abundance of motile fauna were inversely correlated with tidal elevation at north- and south-jetty stations (Figs. 15 and 16). The increase in species richness and abundance at the lower levels is related to the increased structural complexity of the sessile community at those levels. Dean (1981) found a similar relation- ship between structural complexity and the number of motile species on a fouling community in North Carolina. Increased environmental stresses probably also played an important role in limiting the motile epifauna at the upper levels (Connell, 1972). Estimates of species diversity (Appendix H) for the epifaunal assemblages on both jetties paralleled the patterns noted for species number and abundance. However, no discernible trends were observed in these parameters that could be attributed to the effects of differing degrees of exposure (protected versus exposed) or duration of submergence (inner versus outer) of the jetty rocks (Figs. 15 and 16). (2). Community Composition and Vertical Distribution Twenty-three species accounted for approximately 90% of the 7,209 animals collected in suction samples. The remaining 10% (806 animals) were distributed among 108 other species. Thirteen of the 23 numerically dominant species were amphipods, four were isopods, three were molluscs, and three represented other taxa (Fig. 17). With few exceptions, mean densities of these species were greater on the north versus south jetty. This is most likely due to the greater representation of subtidal levels at north-jetty stations, where most of these species were more prevalent. Temporal variations were also observed in the abun- dance of these species on both the north and south jetties (Figs. 18 and 19). 45 Table 8. Number of individuals and number of species of each major taxon of motile macroinvertebrates from the north and south jetties. NORTH JETTY SOUTH JETTY No. No. No. No. TAXON of Individuals of Species of Individuals of Species Total Percent Total Percent Total Percent Total Percent Number of Total Number of Total Number of Total Number of Total Amphipoda 2183 53.6 25 21.0 2313 73.7 24 38.1 Isopoda 742 18.2 6 5.0 498 15.9 6 oS) Mollusca 481 11.8 28 23.5 215 6.8 7 11.1 Polychaeta 242 5.9 32 26.9 58 1.8 12 18.8 Decapoda 209 5.1 17 14.3 26 0.8 7 11.1 Echinodermata 159 3.9 4 3.4 2 0.1 1 1.6 Nemertinea 25 0.6 1 0.8 17 0.5 1 1.6 Nematoda 11 0.3 1 0.8 3 0.1 1 1.6 Pycnogonida 8 0.2 3 2.5 7 0.2 3 4.8 Turbellaria 5 0.1 1 0.8 0 0 0 0 Sipunculida 4 0.1 1 0.8 0 0 0 0 -Mysidacea 0 0 0 0 1 <0.1 1 1.6 TOTAL 4069 119 3140 63 46 yoeo ye eunestde eTT}ow jo setoeds jo zequnu pue souepuNge ey} Jo UOTSSe1Ze1 APOUTT 08 “UOTIBASTO TPT} JO UoTJoOUNZ e se uotT}eIS AQJef YIA0U S31I93dS JO YAISWNN 09 Ov o2 Zvi + *S0'0- =A 16'0- =! 002 I3N 0 IdN ® O3NQ OdN @ os! (249002 Jed ‘ON) ALISN3G NVIW ool Os 091+ xE0'0- =A a p8'O- =4 fo) °CT ernst y (W) 13A37 1VO0IL 47 *UOTIEASTO [Tept} JO uoTJoOuNF e se uoTIeRIS ARO YANOS YoeO We eunejtde o[tqJow Jo sotoods jo Aequnu pue soUepUNge sy} JO UOTSSeAZeA APBUTI “OT 2ANn3Ty (gw2002 40d ‘oNn) S31I03dS JO YSSWNN ALISNAG NVIW 09 Ov 02 002 os! ool os GS'1+%20'0- =A 201 + X10'0- =A 18:0- =3 b8'0- =! = e@ | | i | 4 o > (om I3s o o fh IdS @ < o3s a OdS @ Se 3 48 NORTH JETTY 10 (n=56) oe finn SOUTH JETTY (n= 50) no o [o) wo (zgW9002 48d ON)ALISN3SG NVAW 49 afios adoundooy pyDaD12 D2uDdg ‘ds p2poryduy snxB auo1y) vadeuro xurd7psoun p4pun] sithzey ungpjuepLiponb Duotavydg sumtof117f D1 1aUucsyoUty DADpnys s1e01e0DIDg papqoundispynb D721 2pDing e1nuay 011 atdponibg 3 tus sOquay szeuazmDYy a7 DAydDg Bla7Eqan sOqUulyT ‘ds stsdomuuny pypjnozpuedd 0321en sds g0y20uUa4S 818U92128D1q SNLUOYIYOULY Dug] inbe v17auIdn7 s21aa7 sndowsn7gz p4paypf pssvp sds um21ydoiog siquruad v1, aLdn) J OTHERS MOLLUSCA ISOPODA AMPHIPODA Estimates of overall mean density for the dominant motile macro- Figure 17. Species which contributed greater than 1% of the total number on either jetty are included. invertebrates of both jetties. 25 1979 (n=14) 10 wo 1980 (n= 14) (n=14) 198| wo oO wo wo fo} wo (zWw9002 49d ON) ALISNSGO NVIJW 1982 (n=14) 10 tfivs adoundoay DADO] Oe Eee sds vzpo2yduy snaB auo1y) paueu1a xu1djpsouy] papun}] sqthqsy unzoquapiaponb puoranydg swmiof217f D1 Leuosyordd Dyppnva s1a0daaDIDd pyvzound1zponb v1] apbind sinuaq D1] aadvonItDg UT UB iS OCT, sisuaipmpy azDhyang 1WOqSqaN SOQUAT ‘ds stsdiunuuvy pypyno1puaddy 04211 2W ‘ds 20440Uu27g 818U91118DUq SN2ZUOYZYOTW vaqijinba v11aadnp 81a2a1 sndowsy1g pqvo1pf vssve sds um2zydouog s2qzupued v7] aWNdn9 } MOLLUSCA OTHERS ISOPODA AMPHIPODA Annual changes in the density of dominant motile macroinvertebrates from the north jetty. Figure 18. Estimates represent mean values from all stations during a particular year. 50 SPRING 1980 (n=10) SUMMER 1980 (n 210) 2 E 5 FALL 1980 (n=10) g 2° 2002 48d ON) ALISNAG NV wo o N 1981 oO nN WINTER 10) 1982 (n= 4) (n= SUMMER 1981 a wW = 2 =) no o Oo Wwos = 2fivs adoundooy p4pAv1a DLUDAg Dauaur9 xurd]pseu/) pypun7] s2thzey ungoquaprrponb vuoxsvydg sumtofi17f D1 JaucEYyo1Ag pyvpnvo s1aodaoDsDd nqgpqoundisponb v1] apDubd sinuay D1] audvoDADg CUP Uues SOQUST, qiazeqean soqua] ‘ds s7sdoupummy pgpjna2zpuaddy 0721 2n ‘ds ao0yqouaqs 818Ua1] 18D SNLUOYWYOW” baqzjinba v1. aad09 81027 endowsn7q pypo1pf vesve sds um1ydoaog e1qupuad p11 aXdn9 OTHERS MOLLUSCA ISOPODA AMPHIPODA Seasonal and annual changes in the density of dominant motile macroinvertebrates from the south jetty. Figure 19. stimates E represent mean values from all stations during a particular season. 51 (a). North Jetty Samples collected during the first summer after north- jetty construction contained only 13 of the 23 dominant species, two of which, Paradella quadripunctata and Elasmopus levis, were substantially more abundant than the others (Fig. 18). Paradella quadripunctata reached maximum abundance at the +l-m level and was largely restricted to the intertidal zone (Fig. 20). It has previously been recorded only from Puerto Rico (Bowman, pers. comm.), where it was commonly found on red algae in rocky intertidal environments (Menzies and Glynn, 1968). The other dominant, #. levis, was found at all levels, but it was most abundant at the MLW and -l-m levels. This species is a known inhabitant of rocky intertidal and shallow-water zones of New England and the Gulf of Mexico (Bousfield, 1973; McKinney, 1977). Densities of both P. quadripunctata and HE. levis were greatly reduced by the summer of 1980, and remained low for the rest of the study period (Fig. 18). An increase occurred in the number of species collected during the second year, with 19 of the 23 most abundant species being present. However, the densities of most of these species were lower than in other years, with the exception of the gastropod Astyrts lunata. This mollusk is a common inhabitant of shallow coastal waters of South Carolina (Zingmark, 1978), and was most abundant at subtidal levels (Fig. 20). By 1981, densities of most species had increased and all of the 23 dominants were present on the rocks (Fig. 18). Astyrts lunata remained abundant, but three species of amphipods were even more common. Caprella penantts, morphologically adapted for clinging to fouling biota (Bynum, 1978), was most abundant at the subtidal levels, as were Erichthonius brastltensts and Melitta appendiculata. The latter two species reached maximum densities during 1981 along with the ophiuroid Amphtodtia sp. All but one of the 23 dominant species were still present on the jetty rocks during the last sampling period. Caprella penantts remained the most abundant species and, together with the amphipod Corophium sp, (probably C. ascherusicum) and two species of isopods (P. quadripunctata and Paracerceis caudata), numerically dominated the motile fauna (Fig. 18). The two morphologically similar sphaeromatid isopods, P. quadripunctata and P. caudata, showed considerable niche separation, with the former being found in the intertidal region and the latter being largely restricted to the subtidal zone (Fig. 20). Menzies and Glynn (1968) also reported that P. caudata occurred mainly among algae, seagrass, sponges and the like in subtidal waters. Corophtwn sp. showed significantly greater abundance on the protected side of the north jetty than on the exposed side (p < 0.05, Mann-Whitney U-test), but was the only dominant species to show a significant difference. Cluster analysis of north jetty suction data provides further evidence of annual changes in overall community structure of the motile epifauna (Fig. 21). Collections from the four stations sampled in 1979 grouped Sy MOLLUSCA OTHERS ISOPODA 10 individuals per 200cm2 tH SCALE AMPHIPODA South Jetty a> = fad @ 2 £ =- r o 2 (W) 13A31 2hps adoupdoay pypap1 2 DiuDig sds pipozyduy Dadeu1o rurd7ps0u/) sna auo21y) DADUN] SUlhizsy unzoquepitponb puikaDyds sumrof212f D1 12U0sYoIdg DZDpNDo s1eotaoDIDg pypjoundiurponb pv] 1 appiog sinuaqy 017 aadpopiDg paqziynba v1 12adn9 yams soquey qraqeqan soquay p4pjnorpuaddp 1421 aH p4Do1D{ psspr ezeust]1eDaq anqucysyourg “ds sisdommummy sds umzydatop “ds 20470Ua4S s2aa1 sndowsp2zZ siqupuad p11 andD) sisuaipmy a7Dhydldg oO TF =e Estimates riods combined. ing pe ties from all stations and sampli a Vertical distribution of the dominant motile macroinvertebrates on both jetties. densi represent mean Figure 20. Group Season Station SU 80 NPI SU 82 NPO SU 8l NPO 3 SU 8| NPI SU 8l NEO SU 8 NET SU 82 NEO es) cay SU See NETL SU 82 NPI SU 80 NPO 2 su 80 ~_—SOWiNNECO SU 80 _NET SU 79. ~NPO SU 79 NPL SU 79 NEO SU 79 NEI 8 6 4 ne 0 =e SIMILARITY Figure 21. Normal cluster dendrogram of north jetty suction data indicating station grounvs formed using the Bray-Curtis similarity coefficient. 54 together (Group 1) and were relatively dissimilar to those obtained in later years. For the most part, 1980 samples were also relatively similar to one another (Group 2) and relatively dissimilar to the other groups. Collections from 1981 and 1982 formed Group 3, indicating higher similarity of faunal composition during these later years as compared with earlier years. This suggests that the initial changes in the motile community were more significant than those in later years. Differences in faunal similarity on wave-exposed versus protected sides were not pronounced. (b). South Jetty Four species of amphipods were the only abundant motile fauna on the south jetty during the spring of 1980, shortly after construction was completed. These amphipods were generally restricted to MLW and below (Fig. 20), and their densities were greater than those for other species during any sampling period, with the excep- tion of Paradella quadripunetata (Fig. 19). Dense mats of the hydroid Tubularia crocea occurred during this season (Appendices C and D), supporting very high densities of the caprellids C. penantis and C. equtlibra, and the tubicolous gammarids Jassa falcata and Corophium sp. Bynum (1978) also noted an association between this hydroid and C. penantis in North Carolina waters. Other species found during the spring were very low in abundance (Fig. 19). By the following summer, all of the initially dominant species had declined considerably, in number, and only C. penantits remained at moderate densities. Most of the other species collected during this season were rare, and only 10 of the 23 dominant species were present. Much of this decline may be attributed to decreased coverage by 7. crocea, which was present only when water temperatures were low. Differences in the relative abundance of motile species were also noted between the north and south jetties during this season (Figs. 18 and 19). Dominance continued to change during the fall and winter seasons, with P. quadrtpunetata being most abundant during the fall and J. falcata dominating during winter. Astyris lunata was also relatively common during both seasons, as were a number of amphipod species. Many of these seasonal changes are attributable to the reproductive periodicities of the motile species, and to changes in dominance of sessile taxa (Figs. 9 and 11). During the last two years, the high intertidal isopod P. quadrtpunctata was the most abundant species (Figs. 19 and 20), and it was frequently observed in the empty tests of the barnacle C. fragilis. Shoaling had occurred around the south jetty by this time, resulting in the loss of most lower sampling levels. As a consequence, the decreased abundance of the other dominants (generally most common subtidally) was probably not strictly a function of temporal change, but largely reflected this intertidal sampling bias. The poor representation of subtidal levels on the south jetty also obscured the interpretation of cluster analysis of the suction data from 55 this jetty. No clear patterns of temporal or spatial similarity were observed, and as a result, the cluster dendrogram is not presented here. (3). General Discussion Numerous studies have assessed the sessile component of fouling communities in rocky intertidal and subtidal habitats (see Connell, 1972; Paine, 1974 for reviews), but few have examined the highly motile epifauna associated with those fouling communities (Dean, 1981). This study provided the opportunity to characterize the species composi- tion, distribution, and abundance of the motile macroinvertebrates on both jetties, and to relate these to the development of the sessile community. Similarities were observed in the patterns of temporal change in abundance, dominance, and species composition between the motile epifauna and the sessile biota of the jetties. Both of these biotic components achieved high overall densities relatively quickly, particu- larly on the south jetty. The dense stands of 7. crocea, which generally develop in this geographic area only during periods of low water temperature (Woods Hole Oceanographic Institution, 1952) provided excellent habitat during the spring of 1980 for several species of motile amphipods, including C. penantts. This amphipod is reproductively adapted to rapidly colonizing habitats which undergo frequent reduction in the density of the sessile inhabitats (Bynum, 1978) such as that observed for Tf. crocea. Increased faunal richness at the subtidal levels as compared with the intertidal zone was also observed within both the motile and sessile communities. Dean (1981) noted that the richness of motile species was positively correlated with the richness of the sessile species in the fouling community he investigated. Similarly, the structurally complex community of sessile species, such as ascidians, bryozoans, hydroids, and algae found at subtidal levels on the Murrells Inlet jetties, enhanced the development of a diverse assemblage of motile species. Dean (1981) also described a negative correlation between motile species richness and dominance of the sessile fauna by one or a few species. The over- whelming dominance by Chthamalus fragilts at upper intertidal levels is not conducive to the development of a rich motile epifaunal community at those levels on the jetties investigated in this study, where P. quadripunetata was the only abundant motile species. Certain differences were noted in the development of the sessile and motile communities on the jetties. For example, the number of motile species on the jetties increased over time, but the number of sessile species was relatively constant during the study period. Furthermore, the vertical zonation observed among the sessile species was not as well defined for the motile forms. The distinct bands of sessile organisms are a result of the sedentary nature of these organisms and their compe- tition for space in suitable environmental conditions. The motility of free-living organisms, on the other hand, allows them to migrate over a wide vertical range during periods of submergence. 56 c. Jetty Fishes (1). Species Composition A list of the fish species observed or collected near the jetties is presented in Table 9. Many of the species were found only in the gill nets placed on sand bottom near the jetties, and it is likely that most of those fishes would have been captured even if the jetties were not present. Even so, jetties are known to serve as excellent artificial reefs, increasing both the abundance and diversity of ichthyofauna in areas where they are present (Hastings, 1972, 1978; Hurme, 1979). A large number of fishes were observed on or near the jetty rocks while scuba diving. The two species most frequently seen were the crested blenny (Hypleurochilus geminatus) and black sea bass (Centropristts striata). Both species are commonly associated with reefs (Parker et al., 1979; Powles and Barans, 1980; Middleditch, 1981), and black sea bass are recreationally as well as commercially important in South Carolina waters. The majority of C. strtata seen on the jetty rocks were juveniles, suggesting that this species is utilizing the jetties as a nursery ground. Other recreationally important species frequently encountered around the jetties included sheepshead (Archosargus probatocephalus), Atlantic spadefish (Chaetodipterus faber), spotted seatrout (Cynoseton nebulosus), bluefish (Pomatomus saltatrix), mullet (Mugil sp.) and southern flounder (Paralichthys lethostigma). (2). Food Habits Data on the food habits of selected fishes is presented in Table 10. With the exception of black sea bass which were easily captured, it was somewhat difficult to obtain sufficient specimens for a thorough analysis of diets. Even so, the analysis of fish stomachs indicated that those species of most recreational importance are utilizing the jetty biota as food. Black sea bass consumed primarily decapods and small fish which were common on the rocks. The two major species consumed (by volume) were the small crab Weopanope sayt and the crested blenny (Table 10). Other important food items included amphipod, isopod and polychaete species which were commonly found on the rocks (Appendices F and G). Decapods and small fishes are also important in the diet of black sea bass found in offshore hard bottom reefs of the South Atlantic Bight (Sedberry and Nimmich, In press). Spadefish mainly consumed a different component of the jetty prey community based on stomach content analysis (Table 10). Primary food items appeared to be the sessile, colonial forms such as sponges, hydroids, bryozoans and algae. Amphipods were also important in the diet of spade- fish, but most amphipods eaten were those commonly found on the colonial taxa, such as Caprella spp. Therefore, it is possible that amphipods were only incidentally consumed during feeding on the sessile biota. 57 Table 9. Species of fishes observed on or near the jetty at Murrells Inlet during field studies, 1979-1982. Gill Blackfish Beach Hook & SPECIES Diving Net Trap Seine Line eee Atlantic sharpnose shark (Rhtzoprtonodon terraenovae) + Rays (Family Dasyatidae—species undetermined) + Cownose ray (Rhinoptera bonasus) + Atlantic menhaden (Brevoortia tyrannis) + Threadfin shad (Dorosoma petenense) + Atlantic thread herring (Optsthonoma oglinum) + Striped anchovy (Anchoa hepsetus) + + Oyster toadfish (Opsanus tau) + Black sea bass (Centropristis striata) + + + Bluefish (Pomatomus saltatriz) + + Atlantic bumper (Chloroscombrus chrysurus) + Florida pompano (Trachinotus carolinus) + Lookdown (Selene vomer) + + Pigfish (Orthopristis chrysoptera) + Bluestriped grunt (Haemulon scturus) ot Sheepshead (Archosargus probatocephalus) + + Pinfish (Lagodon rhombotdes) + Spottail pinfish (Diplodus holbrookt) + Spotted seatrout (Cynoscton nebulosus) a3 Spot (Letostomus xanthurus) + + Atlantic spadefish (Chaetodipterus faber) + + Butterflyfish (Chaetodon sp.) oP Damselfish (Pomacentrus sp.) ap Tautog (Tautoga onitis) + + Mullet (MugtZ sp.) 1 Southern stargazer (Astroscopus y-graecum) + Crested blenny (Aypleurochilus gemtnatus) + Feather blenny (Zypsoblennius hentzt) + + Doctorfish (Acanthurus chirurgus) + Atlantic cutlassfish (Trtchiurus lepturus) oe Spanish mackerel (Scomberomorus maculatus) + King mackerel (Scomberomorus cavalla) + Southern flounder (Paralichthys lethostigma) + + Northern puffer (Sphoerotdes maculatus) + Striped burrfish (Chilomycterus schoepft) + 58 8766 96S8 8°7L TIT SE6E 768L 7°€L 9°S IT? 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Nearly 99% of the food items, by volume, were either mussels (Brachtdontes exustus) or algae. As noted previously, the decline in subtidal mussel density on the north jetty was attributed largely to predation. The stomach content analysis supports this hypothesis since B. exustus represented a major component of the sheepshead's diet. Algae, which were the most prevalent taxa on the rocks during the latter part of the study, represented a major component of stomach contents from sheepshead captured in 1981 and 1982. Stomachs from three other species were analyzed even though two of them, Haemulon seturus and Tautoga onttis, are not generally considered to be recreationally important in South Carolina. Stomach contents from the one bluestriped grunt captured on the rock indicated that this species feeds mostly on algae, amphipods and isopods. The only tautog captured had been feeding on isopods, amphipods and decapods found on the rocks. Five southern flounder, P. lethostigma, were also captured for stomach content analysis but all stomachs were empty. Flounder were probably feeding at night around the Murrells Inlet jetties, The stomach content analysis documents that the jetty fauna and flora are important food sources for the recreationally important fishes. Additionally, the results suggest that many of the fishes are minimizing competition for these food resources by concentrating on different components of the community. Similar divergence in food habits has been noted for other communities of sympatric fish species (Ross, 1977). V. SUMMARY AND CONCLUSIONS 1. Rock jetties recently constructed at Murrells Inlet, South Carolina, provided a valuable opportunity to study colonization and community development patterns of biota on rocky substrata. Previous studies of this type have been very limited along the southeastern coast of the United States. 2. Construction began on the Murrells Inlet Navigation Project during the autumn of 1977 in order to provide a stabilized entrance channel to the ocean. The seaward terminus of the north jetty was completed by December 1978, and annual sampling was initiated at four stations on that jetty during the summer of 1979. The seaward terminus of the south jetty was completed by March 1980. Sampling began at four stations on that jetty in the spring of the same year and continued at quarterly intervals for the first year. After that, sampling was restricted to once a year during summer, as on the north jetty. The north jetty was studied over a four-year period, whereas the south jetty was studied for a three-year period. Two of the stations on each jetty were on the wave- exposed side and two others were located on the protected channel side. 3. At each station, sessile biota was assessed at 7 or 8 levels, depending on station location. Intertidal levels were located at 0.5-m intervals from mean low water (MLW) to 2.5 m above MLW; subtidal levels 62 were located at 1-m intervals to a maximum depth of -2.0 m. Sampling at all levels involved both line-transect and photographed-quadrat census techniques. Motile macroinvertebrates were also sampled at +l-m, MLW, -l-m, and -2-m levels at stations where water depths were sufficient to use a quantitative suction sampler. A more limited effort was made to assess fish. Sampling techniques included collections by gill nets, traps, seine net, hook and line and observations by scuba divers. Fish stomachs were also collected for diet analysis, and hydrographic samples were collected during every sampling period. 4. Water temperature, salinity, clarity, and dissolved oxygen reflected the expected hydrographic patterns in the area. Temperatures ranged from 5.89 - 30.3°C, salinity from 34.5 - 36.1 °/oo0, dissolved oxygen was almost always near saturation, and water clarity varied from 0.7 to 2.5 m. 5. Results of the biological assessments indicated that a diverse assemblage of fauna and flora had colonized both jetties. At least 25 algal species, 195 macroinvertebrate species, and 34 fish species were found over the four-year study period. 6. Coverage by sessile biota was generally as great one year after construction as in subsequent years. This was primarily due to the early settling of blue-green algae and barnacles in the intertidal zone and mussels in the subtidal zone. Biota cover was generally higher in the lower intertidal and subtidal zones than in the upper intertidal zone, which represented a more rigorous physical environment. On the wave- exposed side, cover was often greater at higher levels than on the protected side. Other differences between sides were minimal, presumably because wave energy is moderate in this area. The number of sessile taxa found at the stations was as high after one year as in subsequent years. No differences were noted between sides in terms of the number of species, but more species were present subtidally than intertidally on all sides. 7. Community structure of the sessile biota showed both seasonal and yearly variation on the jetties. Variation between sampling periods was less in the intertidal zone where there was distinct banding of blue-green algae and barnacles (Chthamalus fragilis) in the mid- to high-intertidal area, and mussels (Brachtdontes exustus), barnacles (Balanus spp.) and algae (primarily Ulva sp. and Hypnea musciformis) in the lower region. Temporal variability in the subtidal community was much greater, with dominance changing from mussel cover (B. exustus) to algal, bryozoan, hydroid, and ascidian cover, depending on the side of the jetty and the year. No evidence of a stable "climax" community was found by the end of the study period, with the possible exception of the intertidal biota as noted above. Furthermore, results from other studies suggest that a "climax" community is not likely to occur. Differences observed between sampling periods, and between sides of the jetties, were attributed to predation, competition, natural mortality and light penetration (see text for details). Wave action was not considered to be an important factor with regard to differences in species composition between sides. Although species composition was different, study results paralleled findings obtained in other rocky intertidal systems. 63 8. Differences were noted in community structure on the south versus north jetty. These differences are partly attributed to differences in duration of rock submersion, season of rock placement at the study sites, and water depth. 9. Strong vertical gradients were detected in the distribution of most dominant sessile species. Barnacles and blue-green algae were common only in the intertidal zone. Mussels were most abundant from +0.5 mto-1m. Green algae were abundant at MLW and shallow subtidal depths (-1 m), whereas red algae were common at those levels and at deeper depths (-2 m). Hydroids, bryozoans, and ascidians were generally restricted to subtidal areas, but in that zone less pronounced vertical gradients were observed for species in those taxa. 10. Motile macroinvertebrate species quickly colonized the rocks, with abundances as high in the first year as in subsequent years. Generally, individuals of motile species were more widely distributed among levels than noted for the sessile species, although some were limited to the intertidal or subtidal zone only. The abundance and species richness of motile fauna usually increased at the lower levels sampled, and in both zones, the dominant species were generally amphipods or isopods. Differences were noted between years with respect to overall community structure, but these differences were less than those observed for the sessile community. 1l. Fishes appeared to be quickly attracted to the rocks and were present in high numbers one year after construction of the north jetty. Community composition of the fishes included many recreationally important species, such as black sea bass, sheepshead, spadefish and flounder. Analysis of fish stomach contents indicated that jetty fauna and flora were primary food items for sea bass, sheepshead, spadefish, grunt and tautog. Additionally, it appeared that these fishes minimized competition for food through differences in their primary diet component. 64 VI. LITERATURE CITED Boesch, D. F. 1977. 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"Invertebrate Communities Associated with Hard Bottom Habitats in the South Atlantic Bight," Estuarine, Coastal and Shelf Science. Windell, J. T. 1971. ''Food Analysis and Rate of Digestion," IBP Handbook No. 3: Methods for Assessment of Fish Production in Fresh Waters, W. E. Ricker, ed., Blackwell Scientific Publications, London, pp 215- 226. Woods Hole Oceanographic Institution. 1952. "Marine Fouling and Its Prevention,” U. S. Naval Institute, Annapolis, Md. Zingmark, R. G., ed. 1978. "An Annotated Checklist of the Biota of the Coastal Zone of South Carolina," University of South Carolina Press, Columbia, S. C. 69 debe Wyse - Tuas ip a ait eV, PAI ie . 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' ; in al i he : ; ' ats Wi ou” wy are ww 11> 4 , a ‘ ett Vee Ae. f % So a aS APPENDIX H: ESTIMATES OF SPECIES NUMBER, ABUNDANCE, AND DIVERSITY OF MOTILE EPIFAUNA COLLECTED IN SUCTION SAMPLES- NORTH AND SOUTH JETTY STATIONS H1 Appendix H.1 NPO NPO NPO NPO NPI NPI NPI +1m MLW -1m -2m +1n MLW SU80 3.06 4.26 0.89 5.50 NEO NEO NEO NEO NEL NEI NEI +1m MLW -1n +1m MLW -1m SU79 Estimates of species number, abundance, and diversity of motile epifauna collected in suction samples at north jetty stations. SU82 # # i # # i # # 4 # i # # i # # Appendix H.2 SPO +1m SPO MLW SPO -1m SPI +1m SPI MLW SPI -1m SEO +1m SEO MLW Estimates of species number, abundance, and diversity of motile epifauna collected in suction samples at south jetty stations. SP80 ror onw BON Pee CONN WAOUWW N © POH e e e Re [ony PRO SU80 4 26 IAL 0.56 HOH e 8 PN HOH Bore) ers ee NNFESN FOU KRW PUON fF © © © FA80 i 45 0.00 0.00 H3 W181 al 1 0.00 SU81 oOlonrF NO DATA DATA SU82 NO DATA NO DATA DATA # # i # # i # # i # ip al tf # i # # i # # i # # i Appendix H.2 (Continued) SP80 SU80 FA80 WI81 SU81 SU82 SEI +1m 2 2 3 0 2 a 37 0 1.00 0.44 0.36 = NO NO 1.00 0.44 0.23 = DATA DATA 1.44 0.42 0.55 = SEI MLW 6 3 8 10 14 3 29 47 2.35 1.58 2.40 2.09 NO NO 0.91 1.00 0.80 0.63 DATA DATA OO 1.82 2.08 2.34 * values based on two replicates only H4 # mel # # 4 5 } it a