1. SEASONALITY AND SIZE OF BOONEA IMPRESSA POPULATIONS ON RESTORED AND NATURAL OYSTER REEFS: DOES ‘SIZE’ MATTER?  Loren Coen *, Andrew Hollis *, Steve Roth *, and Rachel Kalisperis + *Marine Resources Research Institute, SCDNR + SC Aquarium, Charleston, SC

2. SEASONALITY AND SIZE OF BOONEA IMPRESSA POPULATIONS ON RESTORED AND NATURAL OYSTER REEFS: DOES ‘SIZE’ MATTER?  Loren Coen *, Andrew Hollis *, Steve Roth *, and Rachel Kalisperis + *Marine Resources Research Institute, SCDNR + SC Aquarium, Charleston, SC Van Gundy vs. Yao Ming

3. Emeryville, CA shell mound, originally reported as over 60’ high >350’ in diameter as over 60’ high >350’ in diameterhttp://sscl.berkeley.edu/arf/publications/54.html Shell Ring, North Edisto, SC (from ~500’ elevation, false color IR) (2-3,000 BP, 200’ diameter, 5’ high, 375,000 bu) Oysters Have Been Consumed for 1000s of Years photo by Anne Shaffer Olympia Oyster Shell mounds

4. Subtidal Oysters, Most Well Known Kent, B.W., 1992. Making dead oysters talk: techniques for analyzing oysters from archaeological sites. MD Historical Trust Kent, B.W., 1992. Making dead oysters talk: techniques for analyzing oysters from archaeological sites. MD Historical Trust Oysters “plastic growth” reflected in different shell ‘shapes’: A) On hard gravel A) On hard gravel B) On soft sediments B) On soft sediments C) Crowded on muddy/shell C) Crowded on muddy/shell Large % oysters in clusters dead shell, 5-95%! (Bahr & Lanier 1981, Russo & Saunders 1989) Above Photo from K. Paynter, UMD

5. Land Snails also Oysters & Middens in an Archaeological Context  Since the 1800s, archaeologists have know that intertidal oysters (se US) were an important subsistence resource in prehistoric coastal societies  Observed shells scattered, heaped or mounded, often >10 m middens  However, their importance the subject of great debate (no direct tests)  Some: argue low nutritional value and other assumptions support a lower ‘value’ for oysters, suggesting seasonal movements and therefore an assumed cold weather resource  Summer months Dermo (20 th century phenomenon?), lower nutritional yield (notions stem from modern obs.)  Others: year round resource, not a subsistence item only (e.g., Russo)  Seasonal models cannot use ‘growth checks’ for oysters vs. other intertidal bivalves in SE vs. subtidal NE pop. (Kent 1988, Russo 1991, others), clusters large % dead shell, fast growth (cf., x.s. hard clams)  Focused excavations in oyster middens have found Boonea to be the most abundant gastropod!  Amazingly, most sampling of natural or constructed oyster reefs, regardless of location/type have not quantified Boonea let alone measured them! ASSUMPTION- “If seasonal size patterns for Boonea can be established, the collection period for oysters in middens can also be established” (Russo 1991)

6. “It is imperative that ecologists and archaeologists work more closely together, creating a truly cross-disciplinary alliance that will help to advance both fields.” Briggs et al. 2006: Why ecology needs archaeologists and archaeology needs ecologists. Frontiers Ecol 4:180-188. Kent, B.W., 1992. Making dead oysters talk: techniques for analyzing oysters from archaeological sites. MD Historical Trust

7. Overview of Talk  Sampling a natural reef populations (n = 3 reps/mo.) oBoonea live:dead ratios oMeasured individual live and dead Boonea using digitized images and measuring software oMeasured all live oysters oRecorded other physical data  Re-examined other SC Boonea data from natural and constructed reef populations  Address the validity of Boonea size in archaeological research across the southeastern U.S. as an indicator of harvesting ‘season’ by indigenous peoples  Utility for other current intertidal/subtidal reef restoration efforts  Noting that few if any have quantified or even noted Boonea in their resident sampling efforts! Since January 2006, we have been sampling at monthly intervals (9 analyzed to date) at a natural intertidal oyster reef: Since January 2006, we have been sampling at monthly intervals (9 analyzed to date) at a natural intertidal oyster reef: 

8. Brumbaugh, Beck, Coen, Craig and Hicks, 2006. TNC, 28pp. Developed Design & Monitoring Manuals, Working Groups, Websites Oyster Restoration Working Group http://www.coastal.edu/marine/sgoyster Coen, Wilber, Walters, Hadley, 2006. SC Sea Grant Burrows et al. 2005. Ch. 4, Vol. 2, Restoration Monitoring of Oyster Reefs, NOAA

9. Mussel densities can exceed 1,300 per m 2 Little Understood Reef ‘Players’ Boonea impressa, an ectoparasite of oysters, Dermo vectors, other relevance? SC over-winters over 33% of the eastern race of oyster catchers (forage & breeding sites) 

10. Boonea impressa, 1-6 mm long

11. What We Know in General About Boonea  Ectoparasite of bivalves, including, but not limited to Crassostrea virginica (e.g., 36 spp. incl. Crepidula, Geukensia, Urosalpinx)  Reproduction all year, with peaks spring-summer; spermatophore, small eggs (<240 µm), 20-250 hatch 3-5 days, larval duration <7 days, some debate whether planktotrophic vs. lecithotrophic development  Feeds on hemolymph from mantle or gill with proboscis  One of the mechanisms purported to be a mode of Dermo (P. marinus) dispersal and transmission among oysters, along with water borne transmission  Can depress host growth rates (by stress or parasite load?), but effects termed by most ‘short-term’?

12. Boonea cont.  Can reach densities >100 individuals/oyster, 20-40 more typical for large oysters, 4-5/oyster common for VA-NJ sites  Gregarious behavior, often moving daily (‘density-vague’, Wilson et al. 1988)  Dredged (=subtidal?) samples often yield few individuals  Snails use an attachment thread to help them maintain their position if dislodged, sensitive to vibration  Live approx. one year, peak reproduction May-July; July- Aug., 1 mm or smaller; Fall, >3-5 mm; often gregarious  In FL, recruiting hermit crabs use dead Boonea shells as they settle out of plankton (P. Wilber pers. obs.)  We have an extensive and unique dataset quantifying live:dead Boonea

13. Long-term studies in SC on natural and constructed intertidal oyster reefs have pointed to the ectoparasitic gastropod, Boonea impressa (Pyramidelidae) as an ‘important’ constituent of reef resident faunas Long-term studies in SC on natural and constructed intertidal oyster reefs have pointed to the ectoparasitic gastropod, Boonea impressa (Pyramidelidae) as an ‘important’ constituent of reef resident faunas Previous B. impressa work in SC and elsewhere has focused on: Previous B. impressa work in SC and elsewhere has focused on:  Systematics, host energetic costs, host-to-host (bivalves) transmission of Perkinsus marinus, effects on host growth and filtration rates, and to a lesser extent snail population ecology Since the early 1990s, archaeologists (e.g., M. Russo, Nat. Park Service) have used Boonea to derive prehistoric seasonal oyster foraging patterns from midden collections in the southeastern U.S. based on their presence/modal size. Since the early 1990s, archaeologists (e.g., M. Russo, Nat. Park Service) have used Boonea to derive prehistoric seasonal oyster foraging patterns from midden collections in the southeastern U.S. based on their presence/modal size. Since 1995, we have separated live and dead Boonea, assessed live:dead ratios (% live), but never measured individuals, nor collected monthly samples Since 1995, we have separated live and dead Boonea, assessed live:dead ratios (% live), but never measured individuals, nor collected monthly samples However, we did observe seasonal, reef ‘age’ and tidal height variations However, we did observe seasonal, reef ‘age’ and tidal height variations GOALS: 1) to look at our Boonea information from numerous sites collected since 1990s; 2) to assess the validity of their assumptions using extant populations, given that archaeologists cannot tease apart live vs. dead animals as we can. Background and Study Goals

14. From: H.W. Wells, 1959. Nautlius Followed sizes for 18 mo. in NC Followed sizes for 18 mo. in NC Presumably measured only live animals? July July

15. NC FL Closest fit between extant and midden Boonea collections, spring

16. Xanthidae O. tau G. bosc B. impressa A. heterochaelis 0 200 400 600 800 1000 1200 On-bottom off-bottom Intertidal Past SC Inlet Creek Work on Boonea Abundance Past SC Inlet Creek Work on Boonea Abundance From: R. Kalisperis (Giotta), 1999 Trays = 76 cm x 53 cm x 13 cm, 0.418 m 2, n = 5 June 12, 1997- June 12, 1998 Adj. ~700/m 2, Jan. 1998 Inlet Creek Long-Term Study Site Mean No. Per 0.42 m 2 (+1SE) 

17. Long-Term Sampling Results From Our Two SC ORP Sites Mean Boonea abundance adj./m 2 on natural and experimental (constructed) reefs (n = 9) at our two Charleston study sites over six years Interestingly, densities on six constructed reefs were consistently low for 6 years of sampling, regardless of time of year Interestingly, densities on six constructed reefs were consistently low for 6 years of sampling, regardless of time of year July natural reef densities were consistently low, whereas January natural reef densities were high. July natural reef densities were consistently low, whereas January natural reef densities were high.

18. ACE Basin Constructed Reefs Boonea - Oyster January 2005 Patterns ACE Basin Constructed Reefs Boonea - Oyster January 2005 Patterns Reefs constructed, 6/2000 w/variety of shell Sampled 1/7/05 w/replicate 0.25 m 2 quadrats ~4.5 yr old  % Mean Live Boonea above bars n = sample size

19. Aerial of Fort Johnson Year-Long Study Site Near Our Lab (9 of 13 months to date) Year-Long Study Site Near Our Lab (9 of 13 months to date) Study site near Fort Johnson  N 32 o.74579’, W 79 o.90356’

20. Boonea Sampling, Dog House Creek Three oyster samples (0.125 m 2 /mo.), sieved over 0.5 mm mesh

21. Boonea Monthly Collections (both live and dead) The 3 replicate, 0.125 m 2 samples sorted

22. Photos using a Nikon Coolpix 995 grids on a Nikon SMZ645 dissecting microscope Grid is 2 x 2 cm Prep for B oonea Imaging Prep for B oonea Imaging

23. Boonea Separated into Live & Dead Categories & then Oriented for Image Capture & Measurement

24. Measuring Live and Dead B oonea Measuring Live and Dead B oonea Claudia using SigmaScan Pro Ver. 5 Measured shell lengths (green lines) from apex to abapical end for each individual (up to 500/sample)

25. General Monthly Patterns: % Live, Size General Monthly Patterns: % Live, Size Densities of live Boonea fell off precipitously until July ’06, when new recruits arrive (<2 mm TL) Densities of live Boonea fell off precipitously until July ’06, when new recruits arrive (<2 mm TL) Within two months, new recruits have nearly doubled in size Within two months, new recruits have nearly doubled in size Populations once again reached maximum densities in 2-3 months Populations once again reached maximum densities in 2-3 months  Compared to above Russo (1991) data, very similar for 9 months we have Compared to above Russo (1991) data, very similar for 9 months we have July **

26. Jan July Jan July Sampling Results From Our Two Long-Term Oyster Restoration (ORP) Sites Mean Boonea abundances for natural & constructed reefs (n = 9) at 2 sites over 6 years. Note: for 1995 & 1996, sampling also occurred in March (95), May (95), Sept. (95), Nov. (95), April (96), & Oct. (96) Low July densities relative to the January natural reef numbers perhaps from turnover of new very small recruits? Low July densities relative to the January natural reef numbers perhaps from turnover of new very small recruits? March, Sept. & Nov. 1995, Oct. 96, densities high also March, Sept. & Nov. 1995, Oct. 96, densities high also 

27.  May 05 Two Long-Term Oyster Restoration Sites March & May 2005 & January 2006 41.2%41.5% 83.6% 66.9% 88.8% 46.5% 73.8% Natural Reefs on 3/3/05 and Exp Reefs on 5/4/05; % Live above Mar 05 May 05 Jan 06 May 05 Mar 05 Jan 06 Numbers Mean Boonea per m 2 (+1SE)

28. Dog House Creek, Live Boonea x Oysters All Monthly (9) Samples 5/06 9/06 8/06 7/06 6/06 1/06 3/06 4/06 2/06 Past efforts have found little or no relationship between ‘oyster size’ and ‘Boonea ‘ densities (all 27 samples) Monthly Means for nine samples

29. Dog House Creek, Boonea Size-Freq., April ’06 Combined, Reps 1-3 Mean size of Live Boonea: 4.75 mm Mean size of Dead Boonea: 3.88 mm 86% (range, 76-91%) total individuals live Live/Dead sizes overlap

30. Dog House Creek, New Recruits in July ’06 Combined, Reps 1-3 Mean size of Live Boonea: 1.62 mm Mean size of Dead Boonea: 3.20 mm 74% (range, 58-90%) total individuals live Live/Dead sizes do not overlap

31. Dog House Creek, All Boonea (Live + Dead) Monthly (9) Samples

32. Dog House Modal Data Analyzed (Mike Russo, NPS) July Aug Sept June % Modal Size Range Boonea Size Range (mm) Live Boonea Data, Russo 1991 Near St. Augustine, FL Our % live results suggest that regardless of collection time, relative ‘numbers’ from month to month are comparable Dead Boonea may decrease overall mean size, but the change is minimal in terms of monthly or seasonal modes or means, as rounded by archaeologists Mean size in SC suggest dead Boonea are probably ‘recent’ dead and not dead from an extended period from months to years Mean dead sizes were smaller from Jan. to June, suggesting they were from that year's cohort For June-August, mean dead sizes were greater vs. mean lives sizes, suggesting that dead snails are stragglers from the previous year’s cohort

33. Preliminary Findings  Recruitment occurs during a very short window in July 06  Individual sample % live values ranged from low 37% (June rep. #2) to a high of 95% (Jan. rep. # 2)  Monthly mean (n =3) live %: from low of 40% (June) to high of 90% (Jan.)  Individuals reach max size and population densities in only a few months, post-hatching  Measurements and monthly sampling appear to explain our previous seasonal observations for Boonea densities at our other SC restoration study sites. Encourage others to look at Boonea, even if just live snails!  Although Boonea from midden archaeological sampling efforts cannot distinguish live from dead, our results when analyzed using their approach (modal %, etc.) seem to support their assumptions and results  Signatures for seasons best approach for midden samples vs. using monthly data After 9 (out of 13) months of sampling:

34. Acknowledgements Special thanks for generous input/efforts: Virginia Shervette, Mike Russo, Nancy Hadley, Evan Ward, David Bushek, Julie Pai, Darin Jones, Emma Gerald, Michael Hodges, Julie Nelson, Amanda Powers, Lee Taylor, Claudia Jendron, Donnia Richardson, Mike Yianopoulos & Bill Anderson