Calcium carbonate production on the central west Florida continental shelf

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Calcium carbonate production on the central west Florida continental shelf

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Title:
Calcium carbonate production on the central west Florida continental shelf
Creator:
Tyner, Elizabeth Carlene.
Place of Publication:
Tampa, Florida
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University of South Florida
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English
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viii, 166 leaves : ill. (some col.), maps ; 29 cm.

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Calcium carbonate ( lcsh )
Continental shelf -- Florida -- Gulf Coast ( lcsh )
Dissertations, Academic -- Marine Science -- Masters -- USF ( FTS )

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General Note:
Thesis (M.S.)--University of South Florida, 2003. Includes bibliographical references (leaves 95-105).

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University of South Florida
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Universtity of South Florida
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All applicable rights reserved by the source institution and holding location.
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029728012 ( ALEPH )
53472947 ( OCLC )
F51-00008 ( USFLDC DOI )
f51.8 ( USFLDC Handle )

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Calcium Carbonate Production on the Central West Florida Continental Shelf by Elizabeth Carlene Tyner A thesis submitted in partial fulfillment of the requirements for the degree of Master of Science College of Marine Science University of South Florida Co-Major Professor: Norman J. Blake, Ph.D. Co-Major Professor: Larry J. Doyle, Ph.D. Pamela Hallock Muller, Ph.D. Date of Approval: May 22,2003 Keywords: biogenic sediment production, Holocene sediments, foraminifera, mollusca, sedimentation rate Copyright 2003, Elizabeth Carlene Tyner

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ACKNOWLEDGEMENTS I am appreciative of the United States Geological Survey for initial funding for this research as part of the West-Central Florida Coastal Studies Project. Thanks also go to John Edkins of the USF College of Marine Science Center for Ocean Technology for his development of the production maps. I especially thank my committee, Norman J. Blake, Ph.D., Larry J. Doyle, Ph.D. and Pamela Hallock Muller, Ph.D., for all of their continued assistance, encouragement, confidence and patience with both me and the research. I am honored to have them as both mentor and friend. And, finally, I am ever grateful to my family, Mom, Dad, and Layton, for their constant support.

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TABLE OF CONTENTS LIST OF TABLES LIST OF FIGURES ASBTRACT CHAPTER I INTRODUCTION Definition of Work Goal of Study Value of Study Scope of Work Carbonate Shelves Geographical Setting Study Area Description of Data CHAPTER II LITERATURE REVIEW CHAPTER III FORAMINIFERAL CARBONATE PRODUCTION Foraminifera Methods Results Data Used Categorization of Organisms Calculations Discussion of Results Production Rate Factors Influencing Production Regional Comparison Fate of Carbonate Produced Conclusions CHAPTER N MOLLUSCAN CARBONATE PRODUCTION Mollusca v Vl Vll 1 1 1 1 2 2 2 4 4 11 15 15 16 16 17 17 21 23 23 24 25 27 27 30 30

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Methods Categorization of Organisms Data Used Calculations Chemical Composition Turnover Rates Micromolluscan Calculations Macromolluscan Calculations Discussion ofResults CHAPTER V ECHINODERM CARBONATE PRODUCTION Echinodermata Methods Categorization of Organisms Data Used Calculations Chemical Composition Turnover Rates Echinoderm Calculations Discussion ofResults CHAPTER VI CORAL AND CALCAREOUS ALGAL PRODUCTION Algae Algal Productivity Corals Coral Productivity Chemical Composition Carbonate Productivity CHAPTER VII PRODUCTION BY OTHER ORGANISMS Annelida Porifera Bryzoans Crustacea Fish Benthic Crustaceans Water Column Biomass Unicellular Production CHAPTER VIII FLORIDA MIDDLE GROUND PRODUCTION CHAPTER IX RESULTS OF CARBONATE PRODUCTION RATES FOR THE CENTRAL WEST FLORIDA SHELF CHAPTER X DISCUSSION OF RESULTS Factors Affecting Evaluation ofEstimations 11 32 32 32 33 34 34 35 36 37 41 41 43 43 44 44 45 47 48 49 50 50 53 54 55 56 57 63 63 64 65 65 65 66 66 67 68 73 74 77

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Sampling Technique 77 Organisms Omitted 78 Unknown Factors 79 Evaluation of Assumptions Made 80 Future Research 81 Fate of Carbonate Produced 83 Depositional Environment 83 Sedimentation Rate 85 Sea Level Stand 86 Global Carbon Dioxide Budget 87 Benefit of Estimated Carbonate Production Rates 88 Uses of Calcium Carbonate 90 Production Rate Comparisons 90 Concluding Remarks 93 REFERENCES CITED 95 APPENDICES 106 Appendix 1: List of Stations on the Central West Florida Shelf 107 Appendix 2: Large Benthic Foraminiferal Production 1974 Transect II 109 Appendix 3: Large Benthic Foraminiferal Production 1974 Transect I 110 Appendix 4: Large Benthic Foraminiferal Production Summer 1975 Transects I and II 111 Appendix 5: Large Benthic Foraminiferal Production Summer 1975 Transect III and Averages 112 Appendix 6: Large Benthic Foraminiferal Production Fall1975 Transects I and II 113 Appendix 7: Large Benthic Foraminiferal Production Fall1975 Transect III and Averages 114 Appendix 8: Large Benthic Foraminiferal Production Winter 1976 Transects I and II 115 Appendix 9: Large Benthic Foraminiferal Production Winter 1976 Transects III and Averages 116 Appendix 10: Small Benthic Foraminiferal Production 1974 Transect II 117 Appendix 11: Small Benthic Foraminiferal Production 1974 Transect I 120 Appendix 12: Small Benthic Foraminiferal Production Summer 1975 Transects I and II 123 Appendix 13: Small Benthic Foraminiferal Production Summer 1975 Transect III and Averages 128 Appendix 14: Small Benthic Foraminiferal Production Fall1975 Transects I, II, III and Averages 133 Appendix 15: Small Benthic Foraminiferal Production Winter 1976 Transects I, II, III and Averages 138 Appendix 16: Foraminiferal Production Rates by Season 1975-1976 143 Appendix 17: Foraminiferal Production Rate Summary 144 111

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Appendix 18: Micromolluscan Production and Species List 197 4 Transect II Appendix 19: Micromolluscan Production and Species List 1974 Transect I Appendix 20: Micromollusc Species List 1975-1976 Appendix 21: Micromolluscan Production 1975-1976 Appendix 22: Archived Macromolluscs 1974 Appendix 23 : Archived Dominant Macromolluscs 1975-1976 Appendix 24: Macromolluscan Production 1974 Appendix 25: Macromolluscan Production 1975-1976 Appendix 26: Macromolluscan Production MMS 1992-1994 Appendix 27: Echinoderm Species List Appendix 28: Echinoderm Production 1974 and 1975-1976 Appendix 29 : Echinoderm Production MMS 1992-1994 Appendix 30: Dominant Archived Calcareous Algae Appendix 31: Archived Hard and Soft Corals Appendix 32: Archived Serpulid Polychaetes Appendix 33: Florida Middle Ground Production Appendix 34: Total Carbonate Production on the Central West Florida Shelf I V 145 148 150 151 152 153 154 155 156 157 159 160 161 162 163 164 165

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LIST OF TABLES Table 1 Foraminiferal production rates for the central west Florida shelf 22 Table 2 Foraminiferal production summary by season 24 Table 3 Comparison of foraminiferal carbonate production rates 26 Table 4 Micromolluscan production rates for the central west Florida shelf 36 Table 5 Macromolluscan production rates for the central west Florida shelf 37 Table 6 Molluscan production rates for the central west Florida shelf 38 Table 7 Echinoderm chemical composition 47 Table 8 Echinoderm production rates for the central west Florida shelf 48 Table 9 Echinoderm production summary by depth range 49 Table 10 Summary of algal carbonate production rates 58 Table 11 Summary of coral carbonate production rates 59 Table 12 Coral and algal production rates for the central west Florida shelf 62 Table 13 Carbonate production rates for the Florida Middle Ground 70 Table 14 Sediment constituents modified from Brooks (1981) 71 Table 15 Calcium carbonate production rates 73 Table 16 Carbonate production rates for varied environments 92 v

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LIST OF FIGURES Figure 1. MAFLA Boxcore Stations 1974 Cruise 7 Figure 2. MAFLA Boxcore Stations 1975-1976 Cruises 8 Figure 3. MMS Boxcore Stations 1992-1994 9 Figure 4. All Stations of the Central West Florida Shelf Study Region 10 Figure 5 Stations with Foraminiferal Production Rates 29 Figure 6. Stations with Molluscan Production Rates 40 Figure 7. Stations with Calcium Carbonate Production Rates 75 Vl

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Calcium Carbonate Production on the Central West Florida Continental Shelf Elizabeth C. Tyner ABSTRACT The central west Florida continental shelf is partially covered by an isolated carbonate sand sheet. Biological production by calcium carbonate secreting organisms provides the primary source of sediment to this shallow warm-water system. For the west Florida shelf region between the Florida Middle Ground and Fort Myers, carbonate production rates were estimated for the major taxa of carbonate-producing organisms. Standing crop and biomass values for these organisms were obtained from boxcore samples collected on three separate cruise surveys. Calcium carbonate is produced in this region at a rate on the order of 102 g CaC03 m-2 yr1 with a sedimentation rate of approximately 1 o-1 mm yr-1 The mollusca, the dominant constituent of the sediments, produce 54% ofthe calcium carbonate, 39% by the micromollusca and 15% by the macromollusca. Foraminifera contribute 37% of the total carbonate produced across the shelf. Calcareous algae provide 8% of the production. Echinoderms, coral and cirripedia all contribute less than 1%. Vll

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Substrate plays an integral role in community composition and production rates and is, in tum, shaped by the resident communities. Although the assemblages vary greatly both spatially and temporally, the short-term patchiness contributes to long-term stability of production rates across and along the shelf. Vlll

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CHAPTER I INTRODUCTION Definition of Work Goal of Study The goal of this study is to determine the rate of calcium carbonate production on the central west Florida continental shelf. Calcium carbonate production is defined as the mass of carbonate produced per year expressed in g CaC03 m-2 yr-1 (Hallock, 1981 ). The rate of carbonate production can aid in determining sedimentation rates and sources of deposition along the Florida coastal zone area, as well as predicting the productivity of the region. Production rates also provide tools for interpretation of the fossil record. This study focuses on the production by carbonate-secreting benthic organisms on the central west Florida shelf. Value of Study The oil-rich northern Gulf of Mexico is among the most extensively explored continental margins while the central west Florida shelf is among the least investigated on the U.S. continental margin and what has been examined has been presented in fragmented ways (MAFLA, 1974; Doyle and Sparks, 1980). Estimation ofthe carbonate productivity for this region should be of interest to those managing the shelf, benthic biologists, sedimentologists, paleoceanographers, modelers and those who derive income 1

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and pleasure from its riches (Canals and Ballesteros, 1997). Knowledge of carbonate production rates by benthic communities is important universally as well as locally as production impacts diverse areas from the global carbon cycle to the economy of the west coast of Florida. This synoptic presentation ofthe processes unique to the central west Florida continental shelf offers a first glimpse at the rate of carbonate production. Scope ofWork Carbonate Shelves Modem carbonate shelves are distinguished by their broad, shallow areas often tropical and subtropical in nature; carbonate-dominated sediments ofbiogenic origin; low terrigenous input; material primarily non-reefal or coral in derivation; and created, influenced and altered by a set of processes unique to themselves (MacGinitie and MacGinitie, 1968; Sellwood, 1978; Kennett, 1992; Canals and Ballesteros, 1997). Continental shelves and slopes comprise 10% ofthe Earth's surface with 35% to 70% of the Holocene carbonates deposited on the shelves (MacGinitie, and MacGinitie, 1968). As important as they are, little of the energy transfer across the carbonate margins is known and generally very few details exist of those benthic communities (MAFLA, 1974; Canals and Ballesteros, 1997). Geographical Setting A nearly classic example of a carbonate shelf is the central west Florida continental shelf of the eastern Gulf of Mexico. This wide, shallow shelf of subtropical affinities with smooth slope, little relief or framework and little terrigenous sediment input, is ecologically rich, highly diverse and a relatively unknown scientific entity 2

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(MAFLA, 1975-1976; Gorsline and Swift, 1977; Sellwood, 1978; Socci and Dinkelman, 1979; Doyle and Sparks, 1980; Murray 1991; Kennett, 1992). Three major water masses influence water movement along shelf and across shelf on the intermediate shelf region: the Caribbean-derived Loop Current, the West Florida Estuarine Gyre and the Florida Bay Waters (MAFLA, 1974, 1975-1976; Gorsline and Swift, 1977; Socci and Dinkelman, 1979; Brooks, 1981 ). Circulation varies seasonally and is also affected by storm events with the bottom waters less affected by the seasonal variations (MAFLA, 1974, 1975-1976; Gorsline and Swift, 1977; Doyle and Sparks, 1980; Brooks, 1981; Hopkins et al., 1981; Li and Weisberg, 1999). The mixed layer is evident to 100 m and flow reverses during the year from northward to southward with no net loss of material along the shelf(MAFLA, 1975-1976; Gorsline and Swift, 1977; Doyle and Sparks, 1980; Li and Weisberg, 1999). Although the system is considered to be stable, the processes are sensitive to perturbation (MAFLA, 1975-1976; Li and Weisberg, 1999). The West Florida Sand Sheet is carbonate-sand dominated and the shelf has been cut off from major clastic input since the Jurassic (MAFLA, 1975-1976; Doyle and Sparks, 1980; Blake et al., 1995). During the last low sea-level stand, a mature fine quartz sand band was deposited from the Appalachian Province on the inner shelf and beaches (MAFLA, 1975-1976; Doyle and Sparks, 1980; Blake et al., 1995). The riverine input, 5,295 cfs discharge, carries little suspended load to the central west Florida shelf, merely one one-hundredth of that from the Mississippi region, with new terrigenous sediment input usually trapped in the bays, estuaries and lagoons (MAFLA, 1975-1976; 3

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Blake et al., 1995). The clear water often found in this region may be due in large part to this lack of clay material in the sediments (MAFLA, 1975-1976). Carbonate sedimentation is controlled by bioproduction and the sediment is largely formed near the site of deposition (Bathurst, 1971; Gorsline and Swift, 1977). The major source of new material into the west Florida system is carbonate in composition (Blake et al., 1995). Study Area The surficial sediments of the central west Florida continental shelf offer a unique study of a carbonate sand sheet isolated from clastic input. The area studied extends on the continental west Florida shelf from the Florida Middle Ground area north ofTarpon Springs southward to Fort Myers. In this stable environment in the eastern Gulf of Mexico, no outstanding topographic features have been noted along the gentle sloping shelfbottom; yet some distinct hydro-biological zones have been identified (MAFLA, 1975-1976). This region enjoys higher species diversity and a higher biomass than other areas within the Gulf of Mexico. The modem sediment found in this carbonate environment is ofbiogenic origin and no riverine sediment has come into this region during the present high stand of sea level (MAFLA, 1975-1976). Description of Data The data used were collected from three separate cruises evaluating the shelf. The Baseline Monitoring Studies, Mississippi, Alabama, Florida (MAFLA) Outer Continental Shelf cruises conducted by Bureau of Land Management (BLM) on two series of cruises, one in 1974 and one in 1975-1976, provided benchmark measurements of certain factors 4

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affected by oil and gas exploitation (MAFLA, 1974). Transects were selected to represent the different biotopes ofthe MAFLA region (MAFLA, 1975-1976). In 1974, 65 stations were sampled in five different transects as noted in Figure 1. The 10 stations ofTransect I west ofTampa in 30 to 40 m depths and the 12 stations of Transect II on and surrounding the Florida Middle Ground at 34 to 54 m depths will be discussed. The box cores were sampled in June 1974. The 1975-1976 cruises expanded upon the 1974 cruises to sample stations three times during the year to represent the three biological seasons of the MAFLA area (MAFLA, 1975-1976). Ofthe 45 stations sampled, 18 stations in three transects lie on the central west Florida shelf as seen in Figure 2. Transect I extends westward of the Fort Myers area and included six stations at 11 to 168m depth. Transect II contains no outstanding topographic features on the gentle sloping bottom and has six stations ranging from 19 to 189 m. Transect III sampled stations on or around the Florida Middle Ground from 20 to 176 m. A total of nine stations lie on the Florida Middle Ground and seven stations from the 1974 cruises were replicated in the 1975-1976 cruises. Analysis was performed for the infauna, epifauna, epiflora, chemistry, sediment composition and water column. Most of the data used here were compiled from the replicate box core samples taken at the 40 stations. Macroinfaunal biomass values also are used from surveys conducted in 1994 by the Minerals Management Service (MMS) of the U.S. Department of the Interior. These cruises surveyed in coastal and shallow open ocean sites along the west coast of Florida to assess the effects of seabed mining activities on the benthic organisms (Blake et al., 5

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1995). Four stations off Egmont Site I, three stations from Site II in Sarasota, one site off Manasota Area III, and one west of Longboat in Site N were evaluated and shown in Figure 3. The replicate box cores were taken from 6 and 13m depths. From these three cruises, 48 stations lie on the central west Florida shelf and are shown collectively in Figure 4 (Appendix 1). Data collected from these surveys are used to calculate estimates of carbonate production rates. The data, although not uniformly nor consistently presented, are among the most extensive yet available for the central west Florida continental shelf and are comprehensive in many ways. Box core samples and observations provide a picture of the benthic carbonate-producing community. 6

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Figure 1. t" 1.1AFLA L o 01:e Ar;,a Dou:orfl Stotlona MAFLA Box core Stations 197 4 Cruise o d'i.fo>..e 0 7

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Figure 2. MAFLA Boxcore Stations 1975-1976 Cruises I I 8

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Figure 3. MMS Boxcore Stations 1992-1994 870 86 eso 840 830 820 PENSACOLA v ; JOO FLOR IDA 290 270 25: ocs GULF OF MEXICO REGION EASTERN 9

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Figure 4. All Stations of the Central West Florida Shelf Study Region -85" 00' -84"10' -83" 20' -82" 30' 29" 10' 29"10' 28" 20' i 28" 20' 'I 27" 30' 27" 30' 26" 40' -85" 00' -84 "1 0' -83" 20' -82" 30' 10

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CHAPTER II LITERATURE REVIEW To date, little research has been published quantifying the calcium carbonate production for marine shelf communities. Many carbonate facies have been described yet few data exist regarding the production and accumulation rates or thickness of the Holocene shelf carbonate (Milliman, 1993). By far, still the most extensive study available is that of Smith (1970, 1971) who determined the calcium carbonate budget for the southern California continental borderland. His budget includes calculations for chemical and mechanical as well as biological transfer both into and out of the system. Biological production rates ranged from 1 to 102 g CaC03 m-2 yr-1 within the different shelf environments. Smith concluded that 400 g CaC03 m-2 yr-1 is produced by the shallow macrobenthos. For the temperate California borderland, he concluded that a carbonate budget is an adequate method for determining processes controlling the carbonate content of the marine sediment. Chave et al. (1972) estimated the carbonate production by coral reefs by evaluating hypothetical reef systems. Potential, gross and net production rates were estimated. Comparison to previously published data revealed that the production rates determined from their models showed reasonable similarity to the rates determined from actual reef systems. Their research concluded that coral reef production rates were driven 11

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by coral growth rates with gross production on the order of 104 g CaC03 m-2 yr-1 with a net production of 10 3 g CaC03 m-2 yr-1 retained by the reef. Chave et al. (1972) also determined that the production was dependent upon the type ofreefhabitat identified from among four different habitat models. Hubbard et al. (1990) examined the production of calcium carbonate for a shelf edge reef system located in the U. S. Virgin Islands. Coral production dominated the total reef production rates, providing 93% ofthe 1.2 x 10 3 g CaC03 m-2 yr-1 carbonate produced. They found production greater near the shelfbreak and did not find a direct depth relationship. They estimated 75% retention within the reef fabric. Milliman (1993) presented average carbonate production and accumulation on a global scale. He presented average production values for the shallow-water environments of coral reefs, banks and embayments, and continental shelves, as well as the production for the deep sea. He estimated a carbonate production rate of 60 g CaC03 m-2 yr-1 for carbonate-rich shelves dominated by benthic production. Production for non-carbonate shelves may be as low as 25 g CaC03 m-2 yr-1 ; that of a coral reef complex is 1.5 x 10 3 g CaC03 m-2 yr-1 ; and production may reach a maximum of 3 x 10 3 g CaC03 m-2 yr-1 for production by Halimeda bioherms. Total production of CaC03 worldwide was calculated to be 5 billion tons per year of which 60% is accumulating. Langer et al. (1997) also estimated global ocean carbonate production and the role played by reef foraminifera and found a range of 30 to 1000 g CaC03 m-2 yr-1 They presented a novel method for production calculation by using a numerical transformation to convert skeletal sediment percentages to foraminiferal production rates. They calculated a rate of 1.2 g CaC03 m-2 yr-1 to 120 g CaC03 m-2 yr-1 for foraminiferal 12

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production in low-productivity lagoonal areas and 230 g CaC03 m-2 yr1 for high productivity reefal areas. Light was the key factor in determining production rates for the larger symbiont-bearing foraminifera. They also discussed the role of carbonate production in the global C02 cycle. Kleypas (1997) proposed a model estimating carbonate production for shallow, tropical regions focusing on physical and chemical controls. Kleypas also found light to be the primary control in calcification rates using corals as the dominant carbonate producer. Bosence (1989) reviewed previous works on production and calculated biogenic carbonate production in Florida Bay. His estimates are 10 3 g CaC03 m-2 yr-1 for the Buchanan Banks and 300 g CaC03 m-2 yr-1 for the Upper Cross Bank. Canals and Ballesteros (1997) identified eight different benthic communities on the Mallorca-Menorca shelf in the northwestern Mediterranean Sea and figured carbonate production for each. Most are algal dominated in depths less than 90 m. They found production higher on rocky bottoms than soft substrate. The range of rates was 91 g CaC03 m-2 yr-1 to 124 g CaC03 m-2 yr-1 with a mean estimation of 100 g CaC03 m-2 yr-1 Smith and Kinsey (1976) summarized and discussed calcium carbonate production by coral reefs. They estimate that shallow, seaward coral reefs produce around 4 x 103 g CaC03 m-2 yr-1 and protected areas 8 x 102 g CaC03 m-2 yr-1 The discussion included production in relation to sea level changes. Moore (1972) explored carbonate production by subtropical soft-bottom communities in Biscayne, Florida. From observations, a rate of 4 x 102 g CaC03 m-2 yr-1 13

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was calculated for sublittoral communities and 103 g CaC03 m-2 yr-1 for one intertidal area. Molluscs and echinoderms dominated production. Frankovich and Zieman (1994) examined the seagrass Thalassia and its epibionts in Florida Bay. Production estimates ranged from 2 g CaC03 m-2 yr-1 to 280 g CaC03 m-2 -1 yr. Hallock (1981) estimated production by Pacific benthic foraminifera and summarized, for comparison, the carbonate production rates for a variety of reef carbonate producers and environments. Production by the foraminifera ranged from 1.5 x 102 g CaC03 m -2 yr-1 to 2. 8 x 1 03 g CaC03 m -2 yr-1 Thus, much foundational work on production by carbonate-secreting organisms has been published. Some of this research will be used to aid in estimating rates for the central west Florida shelf. The results obtained by others will be compared to the findings of this study in the discussion. 14

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CHAPTER III FORAMINIFERAL CARBONATE PRODUCTION Foraminifera Foraminifera comprise a group of diverse marine shell-bearing protozoans (Kennett, 1992). Foraminiferal tests are predominately calcitic or agglutinate (Kennett, 1992). The only aragonitic species in this central west Florida shelf study area are a few small benthic species of the Suborder Robertinina; Bulimina, Cassidulina, Discorbis, Hoeg/undina, and Saracenaria (Murray, 1991). Benthic foraminifera dominate in diversity, number and production in shallow warm water, shelf carbonate systems and their production and accumulation rates may be as much as one to three orders of magnitude larger than the planktic production rates (Milliman, 1993). However, planktic foraminifera contribute far more calcium carbonate to the global ocean system as they cover oceanic basin area orders of magnitude greater than the benthic species (Milliman, 1993; Langer et al., 1997). Since the Paleozoic Era, foraminifera have been a common benthic organism in warm, shallow seas, like the Gulf of Mexico (Hallock, 1981 ). Frequently, foraminifera are classified by size, larger foraminifera range from 10 -3 to 10 -2 m and smaller foraminifera from 10 -5 to 10-3m (Murray, 1991; Kennett, 1992). The larger benthic foraminifera generate significant amounts of carbonate sediments and are 15

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key producers of sand-sized carbonate sediments in reefal and shallow water carbonate systems (Hallock, 1981 ). Methods Data Used The data collected and compiled for the MAFLA lease area studies of 1974 and 1975-1976 include very detailed analyses of the living benthic foraminifera on the central west Florida shelf. Due to their small size, the foraminifera are not easily quantified by biomass (wet weight) figures (Murray, 1973) Therefore, Bock (MAFLA, 1974; 19751976) presented the living benthic foraminiferal data collected in 1974 by number and species per sample area. The 1975-1976 data were listed by total foraminiferal density per sample for three distinct seasons and by number and species per sample for the summer. Samples were extracted from 21.3 em x 30.5 em x 15 em box cores taken during these two MAFLA studies. Results are presented from evaluation of samples from a total of 40 sites within this study area. A 2.5 em diameter by 15 em plug was removed from each of two cores The upper 3 em of each sub core were preserved in glutaraldehyde for the identification of living foraminifera by protoplasm content. The remaining sediment was archived for fossil faunal analysis and comparison with the recent past to determine natural changes in the environment (MAFLA, 1974; 1975-1976) The subcore samples were wet-sieved through a 63 Jlm sieve to remove the finer sediment. Three hundred specimens per sample were picked, mounted and identified by the MAFLA researchers. Analysis included identification of planktic to benthic ratio, percentage of living 16

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specimens by species per sample, and number of total and living specimens per milliliter of sediment (MAFLA, 1974; 1975-6). Categorization of Organisms To estimate carbonate production rates from these archival foraminiferal data sets, the species were first divided by test composition, either calcareous or agglutinate. Calcareous taxa were then separated by size into three classes with no distinction between calcitic or aragonitic tests. The large algal symbiont-bearing foraminifera were divided into two groups by suborder: 1) the large rotaliine genera Amphistegina and Gypsina, which secrete hyaline, perforate, calcite tests; and, 2) the milioline taxa Archaias, Peneroplis and Sorites which secrete porcellaneous high Mg calcite tests (Parker, 1982). All other foraminifera were placed into the small-size class. The large, symbiont-bearing foraminifera were again divided by depth of occurrence, with stations in the 11 to 30 m range in one group and the stations in the 31 to 189 m in another. Production rates were estimated for each of these groups. Calculations Annual production of calcium carbonate by benthic foraminifera depends upon four variables: standing crop, proportion of individuals that reproduce, frequency of reproduction, and the number of new individuals produced from each reproductive cycle (Murray, 1967; Muller, 1974). Production also can be estimated using standing crop and frequency of reproduction (Muller, 1974). Hallock (1981) outlined an alternative method to estimate carbonate production by the use of turnover rates. Turnover rate is the portion of the total amount of a substance released in a given length of time (Hallock, 1981 ). Turilover rate can be calculated by dividing the annual sediment production by the 17

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average standing crop. Unfortunately, the annual sediment production by foraminifera is unknown for the central west Florida shelf. For this study, only standing crop values are known for the MAFLA region. Therefore, production rates must be derived from research done by others from other regions to extrapolate the production in this region. For the large rotaliine foraminifera, Hallock (1981) determined carbonate production from detailed growth rate and standing crop data collected at rockpool, reef flat and slope sites in Hawaii and Palau. Production is considerably higher in Palau than Hawaii due to increased growth rates in Palau as well as the production by the family Calcarinidae (Hallock, 1981 ). The production by Amphistegina gibbosa in the Gulf of Mexico most closely resembles that of Amphistegina lessonii on the reef slope in Hawaii. The Hawaii sites are shallower (5 to 20m depth range compared to 11 to 189m on the Florida Shelf), at lower latitude (21 N compared with 26 to 29 N on the Florida shelf), less seasonal and part of a different carbonate system. For these large, algal symbiont-bearing foraminifera, physical energy associated with water motion and light intensity influence growth rate and reproduction as well as controlling calcification rates (Hallock, 1986; Murray, 1991; Kleypas, 1997). Calcification rates can be as much as two to three times higher in light than dark conditions (Murray, 1991; Lea et al., 1995). Benthic foraminifera secrete more calcium carbonate in high-energy environments, which also increases production (Murray, 1991). The decrease in both light intensity and energy of the system associated with the increase in depth and latitude on the central west Florida shelfleads to a subsequent reduction in calcium carbonate production rates of approximately 75% to 90%. 18

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At the stations of depth range 11 to 30 m, the production rate for the large benthic foraminifera is estimated to be 25% of the Hawaiian production. For the deeper stations (31 to 189 m), rates of production are estimated at 10% of those in Hawaii (Hallock Muller personal communication, 1995). Twenty-five percent ofthe production rate of 3.4 X 1 o-3 g CaC03 m-2 yr-1 for an individual Amphistegina in Hawaii (Muller, 1976) leads to a production rate of 8.5 x 104 g CaC03 m-2 yr-1 at sites up to 30 m in depth, and, 10% of that rate is 3.4 x 104 g CaC03 m-2 yr-1 used in calculating production at depths greater than 30 m for similar species on the central west Florida shelf study area. Therefore, with the use of standing crop data for each species at each site, the following calculation was used to estimate carbonate production, P = 8.5 x 104 g CaC03 m-2 yr-1 x N, where P is carbonate production rate and N is the number of specimens of Amphistegina and Gypsina at each site within the 11 to 30 m depth zone. For the same species at stations of depths greater than 30 m, P = 3.4 x 104 g CaC03 m-2 yr-1 x N. Similarly, production rates for Archaias angulatus and Sorites hofkeri, two milioline foraminifera, were evaluated using growth and production rates from data collected in the Florida Keys (Hallock et al., 1986). Again, production rates are assumed to be lower than the Florida Keys values, which is 1.2 x 10-3 g CaC03 m-2 yr-1 per individual, due to a similar reduction in light intensity and physical energy as with that of the Hawaiian rates. Central west Florida shelf production rates are estimated at approximately 25% ofthe rates in the Florida Keys for stations 30m or less and 10% of the production at stations greater than 30 m. Peneroplis carinatus and Peneroplis proteus 19

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production rates were calculated at 10% of those obtained from the same study ofthe Florida Keys. The production rate for Archaias angulatus and Sorites hojkeri is P = 3 x 10-4 g CaC03 m-2 yr-1 x N, for depths 30 m or less. For Archaias angulatus and Sorites hojkeri for depths 30m and greater and the Peneroplis species at all depths, P = 1.2 x 10-4 g CaC03 m-2 yr-1 x N. The larger symbiont-bearing foraminifera in the central west Florida shelf study area range between 0% and 19% of the total carbonate producing foraminifera with average standing crop values of 5.6 x 104 specimens per m-2 The production rate estimates for the larger foraminifera are found in Appendices 2 through 9. Densities of the smaller benthic foraminifera are considerably higher, averaging 1.6 x 106 specimens per m-2 Total carbonate production for the small benthic foraminifera is estimated by multiplying the observed standing crop by an average production rate derived from that of other warm, shallow, marine carbonate systems. Muller (1976) determined a carbonate production rate of 8.5 x 10-5 g CaC03 m-2 yr-1 for small benthic foraminifera at several sites in Hawaii. Wefer and Lutze (1978) calculated a production rate of 1.2 x 10-5 g CaC03 m -2 yr-1 for small benthic species at depth of26 to 28m in the Baltic Sea. This value, averaged with 25% of the rate from Hawaii, gives a carbonate production rate for the small benthic foraminifera of 1.7 x 10-5 g CaC03 m-2 yr-1 (Muller, 1976; Wefer and Lutze, 1978; Murray, 1991). 20

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0.5 {(8.5 x 10-5 g CaC03 m-2 yr-1 x .25) + (1.2 x 10-5 g CaC03 m-2 yr-1)} = 1 7 x 10-5 g CaC03 m-2 yr-1 for small foraminifera on the central West Florida shelf. This production rate multiplied by the number of small foraminifera per m2 per site estimates the carbonate contribution of the small foraminifera. The production estimates for the small benthic foraminifera are found in Appendices 10 through 15. Langer et al. (1997) presented an alternative method for calculating carbonate production rates of benthic reef foraminifera. They developed a simple mathematical model converting the foraminiferal skeletal sediment component into a production rate for the foraminifera The skeletal sediment component is multiplied by an average value of foraminiferal carbonate production. This upper limit of productivity ranges from a low of 1.2 g m-2 yr -1 for areas of low productivity to a maximum of 6 g m-2 yr -1 in a modem high-productivity reef area. They found that the larger symbiont-bearing foraminifera produce nearly 80% ofthe total carbonate Langer et al. (1997) assumes that all skeletal components are carbonate. Results For the 1975-1976 MAFLA study, the production rates were calculated for each station for a thr ee -season period and a summary of seasonal rates may be found in Appendix 16. Appendix 17lists production for each station. A summary of the total foraminiferal production rates by transect can be found in Table 1. 21

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Table 1 Foraminiferal production rates for the central west Florida shelf 1974 II 1974 I 1975-1976 I 1975-1976 II 1975-1976 III Florida Middle Ground Large Production Range Average 0-48 17 0-86 22 1-33 12 0-12 5 2-8 5 2-22 10 Average for the shelf 0-86 14 Small Production Range Average 4-86 21 6-86 44 7-30 15 7-52 30 6-72 27 4-31 14 4-86 28 Total Production Range Average 12-86 38 6-124 66 16-63 27 14-63 34 9-77 31 9-53 24 6-124 42 Langer et al. (1997) Method Shelf Florida Middle Ground Average for the shelf 12-60 4-21 15 5 13 The average foraminiferal skeletal component of the sediments in the MAFLA region is 3.4% in the Florida Middle Ground and 10% on the surrounding west Florida shelf(Brooks, 1981). Using the Langer et al. (1997) equation, the foraminiferal production could range from 4 to 21 g CaC03 m-2 yr-1 on the Florida Middle Ground and from 12 to 60 g CaC03 m-2 yr-1 on the surrounding shelf. Ifwe continue to consider shelf production as approximately 25% of that for a reef area, the Florida Middle Ground foraminiferal carbonate productivity would be 5 g CaC03 m -2 yr-1 for the sediment constituent model compared with an average of24 g CaC03 m-2 yr-1 from the above calculations. The surrounding shelf area would then have foraminiferal carbonate productivity of 15 g CaC03 m-2 yr-1 compared with the above shelf average of 42 g CaC03 m 2 yr1 The average for the region using this first-order skeletal-component 22

PAGE 33

production estimate would then be 13 g CaC03 m-2 yr"1 compared with the 42 g CaC03 m-2 yr-1 previously calculated. Discussion of Results Production Rate Using this new production model, the average standing crop of the foraminifera in this region ( 1 7 x 106 organisms per square meter of sea floor) produces calcium carbonate at estimated rates of 13 to 42 g CaC03 per square meter per year. The large, benthic, symbiont-bearing foraminifera, only 5% of the total living carbonate producing foraminifera, contribute an average of 32% of the calcium carbonate produced within the system. The small benthic foraminifera, in turn, produce 68% of the calcium carbonate yet make up nearly 95% ofthe assemblage. An average of two distinct species of large foraminifera per station, 7% of the total number of species, adds nearly one third of the total carbonate produced. Muller (1976) notes that three species of larger foraminifera comprising only 44% of the standing crop produced 97% of the carbonate in the nearshore zone of Oahu, Hawaii. Production does vary by station and season. The production estimations range from 6 to 124 g CaC03 m-2 yr-1 across the shelf. The foraminifera live in microhabitats with conditions that vary nominally between stations creating a mosaic of successful patches (MAFLA, 1974; Murray, 1991; Parker, 1982). Standing crop values ranged from 2.6 x 105 carbonate-producing specimens per square meter to 4.5 x 106 specimens per square meter. Seasonal influences like changes in temperature and weather create affects on carbonate production across the shelf, as do the changes in habitat (Bock, 1974; Murray, 1991 ). For the 1975-1976 cruises, highest production estimates were found 23

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during the 1975 fall sampling period with an average of 44 g CaC03 m 2 yr1 Summer 1975 estimates averaged 30 g CaC03 m2 yr1 summer of 1974 averaged 51 g CaC03 m2 yr1 with winter of 1976 having the lowest average of 19 g Ca C03 m-2 yr1 The seasonal variations are listed by station in Appendix 16 and are summarized in Table 2. Table 2 Foraminiferal production summary by seasoning Ca C03 m2 yr1 Summer 1974 Summer 1975 Fall1975 Winter 1976 Average Small Foraminifera Large Foraminifera Total 32 19 51 22 7 30 35 10 45 14 5 19 Individual station estimates ranged from a low of 1 g Ca C03 m2 yr-1 in winter 1976 to a high of 124 g Ca C03 m2 yr1 in the fall of 1975. Factors Influencing Production 24 7 31 A combination ofbiotic and abiotic factors controls the density, assemblage and production rates of the foraminifera on the central west Florida shelf. Key biotic factors include the availability of food, predation and intraand inter-specific competition (Murray, 1973, 1991; Hallock, 1981; Bishof, 1982). Although these factors play a vital role in shaping the benthic community, they are not likely limited by these biotic factors (Murray, 1991). In a relatively stable environment such as this study area, the standing crop does not vary greatly unless a marked change in the food supply occurs (Murray, 1967). Most of the foraminifera have developed non-competitive feeding strategies and herbivores, detritivores, omnivores, and passive suspension feeders share the same environment (Murray, 1991) Even the symbiont-bearing foraminifera actively feed, and the majority of carbon gain is from this feeding (Hallock et al., 1986; Murray, 1991 ). 24

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A greater role in shaping communities and production rates is played by the abiotic factors oftemperature, salinity, dissolved oxygen, pH, depth, physical energy, turbidity, terrigenous input and the nature ofthe substrate (Murray, 1973; Hallock, 1981; Babashoff, 1982; Murray, 1991). Ofthese, salinity, pH and DO are fairly constant across the shelf and very little terrigenous material is input into this region. An expected correlation with depth does not always occur (Murray, 1991). The influence of depth on distribution and standing crop is an indirect one as changes in depth are linked to more important factors such as light and energy. Calculating the different depth zone divisions for the large foraminifera at different carbonate production rates allows for compensation for changes in production that occur with increasing depth. Babashoff (1982) found the primary factor in controlling foraminiferal distribution on the shelf is the texture and content of the surface sediment that an evaluation ofthis data supports. A definite correlation exists between the nature of the substrate and carbonate production. An inverse relationship between grain size and carbonate production occurs over the shelf region (Fig. 4). The production rate model presented here does support the theory that the abiotic physical-chemical setting (namely the nature of substrate, light and physical energy) rather than the biological composition is the most important factor influencing the calcification rate in this marine environment (Smith, 1970; Bock, 1974; Hallock, 1981; Babashof, 1982; Murray 1991; Kennett, 1992; Jayarahu andReddi, 1995). Regional Comparison Despite the abundance and importance of foraminifera to global oceanic carbonate production, little is known of foraminiferal production for much ofthe marine environment, especially on the central west Florida shelf (Parker, 1982; Murray, 1991; 25

PAGE 36

Milliman, 1993; Langer et al., 1997). A comparison of carbonate production rates for some selected foraminiferal species and assemblages is found in Table 3. Table 3 Comparison of foraminiferal carbonate production rates Carbonate Production Rate Habitat Depth Reference in g CaC03 m-2 yr-1 inm 17 S. California 100-1000 Smith (1971) 104 Potential Hypothetical Coral Reef Chave et al. (1972) 101-102 Gross Hypothetical Coral Reef Chave et al. (1972) 156 Hawaii < 10 Muller (1976) 3.1 (Small only) Baltic Sea 27-28 Wefer and Lutze (1978) 50-600 (Large only) Palau 5-20 Hallock (1981) 60 (Archaias) Florida Keys 1 Hallock et al. (1986) 36-130 Florida Reef Langer et al. ( 1997) 8.4 Florida Lagoon Langer et al. (1997) 13-42 W. Florida Shelf 11-189 This study The species assemblage found on the central west Florida shelf closely resembles that outlined by Murray (1991) for an inner shelf zone with a depth range of 0 to 100 m. A greater diversity of species and increased percentage of smaller species is noted on the shelf than at some shallower, reef systems. The shelf area has a high standing crop average of 1. 7 x 106 live benthic specimens per square meter with the majority (95%) being small. Larger foraminifera typically dominate in shallow, reef environments and production should be greater than on the shelf due to this increase in numbers of large species, as well as in their increased calcification rates where light and energy conditions are more favorable. This study's estimate of 13 to 42 g CaC03 m-2 yr -1 as expected, is at the low end of the range of production rates for tropical reef systems 26

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Fate of Carbonate Produced The foraminifera are not the dominant constituent of the sediment on the central west Florida shelf. Brooks (1981) presented the major sediment contributors for the region with the foraminiferal component 3.4% of the sediment at the Florida Middle Grounds and 1 0% on the surrounding central west Florida shelf. The carbonate content of the sediment at the Florida Middle Ground resembles that of other continental shelves and is typical of other deep water reef environments (Brooks, 1981 ). The rate of deposition in the sediment is different at different locations (MAFLA, 1974). Certainly, the small size and thin, fragile walls of these organisms enhance the opportunity for breakdown, reworking and recycling back into the system as well as making it difficult to recover the tests and identify all in the sediments. Larger foraminifera are less likely to be sorted from the sediment (Muller, 1974). The smaller foraminifera or fragments of dead foraminifera ( 1 0-4 m diameter or less) can be transported from the system in suspended or bed load (Murray, 1991) Biological activity, breakdown of the fenestrated tests after death, and destruction by burrowers, borers, grazers, browsers and predators all greatly impact the fate ofthe foraminiferal tests (Brooks, 1981) The total sedimentation rate for this region is low and the foraminifera may contribute as little as 1 0"2 mm per year. An accurate sedimentation rate and constituent analysis would help in defining the fate of the foraminiferal carbonate produced. Conclusions The calcium carbonate production rate for foraminifera on the central west Florida shelf is estimated at 13 to 42 g Ca C03 m-2 yr-1 The large foraminifera comprise 5% of the standing crop and produce 32% of the foraminiferal carbonate. Greater 27

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production occurs during the fall season. Production increases with decreasing sediment grain size. A calcium carbonate production rate estimate for the foraminifera is invaluable in understanding the processes on the central west Florida shelf as well as a useful tool for evaluating such studies as productivity, ecology, life history, interpretation of the fossil record, paleobiology, paleoenvironment, and sedimentation rate. Foraminiferal production rates by station are shown in Figure 5. 28

PAGE 39

F i gure 5 \;; lt. i rll ,,. lotrt II ill I 1'1'1 ru:-1t1111 TIM.I :::t.r 1r. ' I \ Gl I I I 1, ... ,. .. :,. Ita: .. ::\ Sta t ions with Foram i nifera l Production Rates I ' ..... I : ;.J" 29 \ I \ \. \ 0 I \ t; I 1 \ 1"". 1 .. \. ' ::a !' ' g a-2 yr-1 Foraminifera .... 6-10 1' .; II) ::t .... 11 -14 .... 15-19 20-26 .... 27-35 36-39 40-53 -65 .... 66-92 .... 93-124 c .... "" .. .. .,. . ...... :10. ...... .+. w 1:! / A .. . .. \ l i ,_.,:''1 .. ..

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CHAPTER IV MOLLUSCAN CARBONATE PRODUCTION Mollusca The Mollusca are among the best-known groups of marine invertebrates found as living members and in the fossil record (Vinogradov, 1953; Brusca and Brusca, 1990). Although extensively studied since Aristotle, much of their life history and composition is poorly known, especially for the smaller species, complicating the efforts to calculate carbonate production (Vinogradov, 1953; MacGinitie and MacGinitie, 1968; Brusca and Brusca, 1990; Moore, personal communication, 1995). With standing crop values, average turnover rates, and chemical composition, annual carbonate production rates can be estimated. The MAFLA report of 1976 determined that the benthic molluscs in the MAFLA region are patchy in distribution. Variations in species composition and abundance are influenced by a variety of factors including season, sediment type, depth, geography, year, and temperature. Populations are affected seasonally by recruitment and variation in reproductive cycles and larval settlement induced by the Loop Current. Across the central west Florida shelf, the observed increase in density of molluscs in shallower water may indicate an increase in the availability of food. In general for this area, abiotic 30

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factors exert greater influence on these populations than the biotic conditions. The report concluded that the assemblages in the region are unique in both space and time. The live molluscs identified in the MAFLA region belong to the classes Gastropoda, Bivalvia, Scaphopoda, a few Polyplacophora and one Aplacophoran. The mantle of these molluscs secretes a hard, calcareous skeleton as an internal or external shell (Brusca and Brusca, 1990). The gastropods, known familiarly as the snails and slugs, feed by herbivory, predation, parasitism, suspension feeding and browsing and are rare in fine sediment and in depths over 50 meters (MAFLA, 1975-1976; Brusca and Brusca, 1990). Bivalves, distinguished by two hinged shells, are microphagous or filter feeders and suspension feeders (MAFLA, 1975-1976). Scaphopoda (tusk shells) and the Polyplacophora (chitons) are less widely distributed in the benthic fauna of the region (MAFLA, 1975-1976; Brusca and Brusca, 1990). Filter feeding bivalves and browsing gastropods occur nearly equally in the stations sampled. Once settled, these organisms may not move extensively and locomotion can primarily be a benefit in their search for food (MAFLA, 1976). Molluscs often contribute the largest biomass to carbonate rich-systems (Moore et al., 1968). Although the study area is impoverished in live specimens, the samples are rich in dead molluscan material. The shells and fragments of shells of the molluscs comprise the dominant constituent ofthe sediment in this study area: 30% ofthe skeletal fragments in the Florida Middle Ground and 4 7% of skeletal fragments of the surrounding west Florida shelf (Brooks, 1981 ). 31

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Methods Categorization of Organisms Molluscs of the MAFLA region are divided into two classes by size. Micromolluscs are small molluscs, not exceeding 7 mm in size. No distinct taxonomic division separates micromolluscs from the macromolluscs, although some species never exceed this size (Fretter, 1948; MAFLA, 1974). Chave et al. (1972) also classified micromolluscs as 1o-6m2 or approximately 1 mm diameter. Of the 305 total species of molluscs in the entire MAFLA region, 23 species are exclusively micromolluscan, 259 are exclusively macromolluscan and 23 species are common to both. Data Used Live samples of molluscs were collected and identified from the MAFLA region in 1974, in a three-season study in 1975-1976, and in a two-year MMS study of macroinfauna in 1992-1994 (MAFLA, 1974, 1975-1976; Blake et al., 1995). The mollusca were sampled by box core. Live micromolluscs were obtained from a sediment sample from each box core. For the 1974 cruise, the sample tube measured 3 em diameter by 15 em deep, creating a sample size of 7.1 x 10-4 m2 per station (MAFLA, 1974). The MAFLA cruises in 1975-1976 used a 5.5 em diameter tube for sampling and the two samples per station represent an area of 4.75 x 10-3m2 (MAFLA, 1975-1976). These small subsamples may be of inadequate size to represent the live micromollusc assemblage, as the samples were very poor in live specimens (MAFLA, 1974, 19751976). Micromolluscs species lists and density values (number of specimens per square meter) for live specimens are presented for both cruises in Appendices 18 to 21. 32

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Nine replicate box cores per station were used to determine the macromolluscan biomass at each site. The macromolluscan wet weight biomass is expressed as grams per square meter. The 1974 MAFLA cruise biomass represents 0.48 m2 of sea floor, the 1975-1976 MAFLA cruise 0.54 m2 and the MMS cruises of 1992-1994 0.57 m2 Due to variations in bottom composition, the total penetration of each of the box cores changes between stations. Gross biomass data could be affected by these differences as well as by the chance recovery oflarge molluscs (MAFLA, 1974). Composition of the substrate also may limit the use of the box core. Species lists are available for the macromolluscs identified on the central west Florida shelf (Appendix 22 and 23). Calculations Carbonate production rates may be estimated using some combinations of standing crop values, biomass figures, growth rates, turnover rates and chemical composition. Growth rates of the molluscan species display considerable variation and fluctuate widely (Coe and Fitch, 1950). Factors affecting growth rates include age, size, sex, developmental or reproductive stage, annual changes, densities, temperature, species type, availability of food, feeding strategy, geography, sediment type, seasonal changes, year class, environmental competition and environmental conditions (Fretter, 1948; Coe and Fitch, 1950; Comfort, 1957; Wilbur and Owen, 1964; Frank, 1965, 1969; Moore and Lopez, 1975; Stevely, 1978; Vermeij, 1980). Shell growth rates are not continuous; they may be periodic, episodic, determinate and also negative in periods of stress to the organism (Frank, 1969; Milliman, 1974; Moore, 1975; Vermeij, 1980). The relative growth rates may even vary among the different parts of a single organism (Wilbur and Owen, 1964). Little has been published on molluscan shell production and the variable 33

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growth rates for all molluscs make comparison between species or areas difficult (Comfort, 1957; Craig, 1967; Frank, 1969; Bosence, 1989). This inability to quantify the growth rates for most areas precludes the use of growth rate as a factor for carbonate production rate calculation. Chemical Composition The chemical composition of molluscs varies widely by species as well as size and age ofthe organism (Vinogradov, 1953; Milliman, 1974; Hammen, 1980). Little has been published on the exact chemical compositions and discrepancies exist between the methods of analysis and data presentation. Determinations of water content and weight and composition ofthe shell can give only average values (Vinogradov, 1953). Vinogradov (1953) calculated the composition of many molluscs by species. Molluscan shells are nearly exclusively CaC03 (Vinogradov, 1953; Hyman, 1967). Using Vinogradov's (1953) values for species collected in our study area, bivalve shells average 98.2% CaC03 of the shell weight and shell weight is 98.7% CaC03 for the gastropods identified. Therefore, an average of 98.5% of total shell weight as calcium carbonate will be used. Turnover Rates Turnover rates are also highly variable and can be affected by seasonal changes, extent of predation, time to reach maximum size, and the maximum and average life span of an organism (Fretter, 1948; Moore et al., 1968). For many mature communities, turnover rates tend to be low, longevity is high and the ratio of productivity to biomass is also low (Frank, 1969). The micromolluscs have faster turnover rates than the larger molluscs due in part to the effects of predation and that maximum size is more quickly 34

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reached. Average turnover can range from one to ten per year (Fretter, 1948; Chave et al., 1972). Macromolluscs are longer lived and, although some gastropods can reach more than 20 years of age and some pectinids have a lifespan of only one year, a more realistic range would be a three to five year lifespan, with a turnover rate of 1 yr -1 (Coe and Fitch, 1950; Frank, 1969; Chave et al., 1972; Stevely, 1978). The 1974, 1975-1976 and MMS cruise surveys sampled the molluscs by box core and data were presented by density or biomass. From the biomass or density values and estimated turnover rates, production rates can be calculated for micromolluscs and macromolluscs. Micromolluscan Calculations Carbonate production rates were calculated using micromollusc density data from the 1974 and 1975-1976 cruises. The data were first converted from density in sample to specimens per square meter. Live micromolluscan densities range from 0 to 7.8 x 10 3 specimens per square meter. The 1974 cruises found an average of 1.6 x 10 3 specimens m-2 and, a 7.8 x 102 specimens m-2 average was found on the 1975-1976 cruises. Chave et al. (1972) assumed 10 3 g CaC03 per 1 mm diameter organism The micromolluscs in the MAFLA study range in size from 0.25 mm to 7 mm diameter, with a mean size of 3.6 mm. Thus mass is estimated as follows: Shell Mass = 10 3 g CaC03 mm-1 x 3.6 mm specimen = 3.6 x 10 3 g CaC03 per specimen Total calcium carbonate per standing crop may then be estimated from the standing crop by: Calcimass = No of specimens m -2 x (3. 6 x 10 3 g CaC03 per specimen). 35

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An average turnover rate of 10 yr"1 was used to calculate the production rate (Chave, 1972) The results for the production rates are found in Table 4. Table 4 Micromolluscan production rates for the central west Florida shelf in g CaC03 m-2 yr-1 Range Average 197 4 Transect II 20-84 53 197 4 Transect I 20-103 62 197 4 Average 20-103 58 1975-1976 Transect I 8-38 17 1975-1976 Transect II 0 97 28 1975-1976 Transect Ill 5 130 39 1975-1976 Average 0-130 28 Florida Middle Ground 19-84 53 Shelf without Florida Middle Ground 0 130 37 Average for shelf 0-130 40 Macromolluscan Calculations Carbonate production rates for the macromolluscs of the MAFLA region were calculated from wet weight biomass values. Macromolluscan biomass ranges from 0.4 g m -2 to 630 g m-2 across the shelf. Weights ofthe molluscan shells as percent of total wet weight range from over 50% to up to 73% of wet weight biomass (Hammen, 1980). This was confirmed by observations with local species ofthe bivalves Donax, Mercenaria, and Mytilus and the gastropod, Melon gena An average shell weight of 63% of the total molluscan wet weight will be used for the calculations. An average amount of calcium carbonate per shell is 98 5% (Vinogradov 1953) Therefore, 62% of the total biomass is assumed to be calcium carbonate. An average life span for the molluscs of these transects is three to five years for those that survive past the juvenile (Frank 1969; Smith, 36

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1971; Stevely, 1978). A turnover rate of 1 yr"1 for the macromolluscs will be used for estimation. Calcium carbonate content of the molluscs can be estimated from biomass figures by: Mass ofCaC03 ofMacromolluscs (g CaC03 m"2 ) =Molluscan biomass (g wet weight m"2 ) x 63% Mass of shell of Total Weight x 98.5% CaC03 mass. The production rate estimates for the macromolluscs are found in Table 5. Table 5 Macromolluscan production rates for the central west Florida shelf Range Average 197 4 Transect II 0.2 26 6 197 4 Transect I 0.14 1 197 4 Average 0.1-26 4 1975-1976 Transect I 0.3-109 19 1975-1976 Transect II 0.5 5 2 1975-1976 Average 0.3 109 11 MMS Site I 2.5-5 4 MMS Site II 15-46 36 MMS Site III 3 390 132 MMS SiteN 0.5 2.4 1 MMS Site Average 0.5 390 58 Average for shelf 0.1-390 17 Discussion of Results Molluscan carbonate production rates show some variability across the central west Florida shelf. The range in estimated production rates is 0.1 g CaC03 m-2 yr-1 to 390 g CaC03 m-2 yr-1 The total production rate for the 0 to 60 m depth range and the Florida Middle Ground shows a remarkable similarity of71 g CaC03 m -2 yr -1 yet with differences 37

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in the dominant contributor. The micromolluscs show an increase in productivity at depths greater than 20 m while the macromollusc show a decline. The combined rate averages are compiled in Table 6. Table 6 Molluscan production rates for the central west Florida shelf In g CaC03 m2 yr1 Macromollusc Micromollusc Shelf Depth Range Average Average Average 6-20m 53 18 71 29-54 m 3 52 55 90-189m 0.4 6 6 Florida Middle Ground 8 53 61 Area Average 16 40 56 Molluscs of temperate, tropical and subtropical affinities have been identified on the shelf (MAFLA, 1975-1976) Although the biotic and abiotic conditions will affect each species type differently, some trends have been noted for both the micromolluscs and the macromolluscs across the shelf. The molluscs do show some variability with season, depth and bottom type (MAFLA, 1974,1975-1976). Micromolluscan production rates are up to 2.5 times higher in winter than in the fall and summer, which are remarkably similar. The macromolluscs as well show increased production in the winter especially among juveniles (MAFLA 1975-1976) These observations may be the result of a reduced predation pressure in the winter and the likelihood of fall and winter recruitment for the molluscan species (MAFLA, 1974, 1975 1976). Carbonate production rates also increase for all molluscs with an increase in firm substrate and with coarse sand bottom types (MAFLA, 1974, 1975-1976) Trends in 38

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depth also are noted which may be related to the change from fine quartz sand on the inner shelf to the increase in coarse carbonate sand at greater depth or a change in food supply or predation. Macromolluscan biomass is higher on the Florida Middle Ground than the surrounding shelf as expected with the high relief, increase in hard substrate and the tropical nature of the assemblage. Estimated production rates vary between 0.1 g CaC03 m-2 yr-1 and 390 g CaC03 m-2 yr-1 across the shelf. The rates are graphically depicted in Figure 6. The question must be asked whether this three order of magnitude range is truly that great or if molluscs are underrepresented in the samples. Certainly, the limitations ofthe box core allow for larger specimens to have been missed and penetration is less for the harder substrates found on the shelf. The macromolluscs are not all sessile and this mobility may contribute to the variation in samples. Molluscan distribution is patchy on the shelf, species saturation was never reached at some stations; and the patchiness could contribute to an over or an under estimation. The chance recovery of large specimens also would skew the biomass numbers. The micromolluscan samples were extremely impoverished in live specimens and it was noted that some live micromolluscs were missed in the sample processing (MAFLA, 1974, 1975-1976). Molluscan production rates are significant on the central west Florida shelf. For live material, the micromolluscs are second to the foraminifera in abundance and produce carbonate at a rate similar to the low estimate of the foraminiferal production rate. Molluscan skeletal fragments dominate the sediments (Brooks, 1981 ). 39

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Figure 6 \;;11:.1 r.: .fit ,. iu It'! ... f.an:lard Tln.o ::::,r I \ ()I ...... I -141 I, -I._, ... : .. r .... 1 :---t _. Stations with Molluscan Production Rates I ' 14 . : : ;.:'1 12 () ::.-... . 1) \ ; : -. .>r I \ \ \ I \ .. \ \ ,_:...; \::< _,_,,_::--::. 40 '" ... \ ; I ...... ; It ,_,.'', g CaCOJ m-2 yr-1 Mollusca ..... 0 ..... 1 -11 ..... 12-23 ..... 24-35 36-44 45-60 61 -74 5 103 ..... 104 137 ..... 138 408 ... ... ol -17 ..

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CHAPTERV ECHINODERM CARBONATE PRODUCTION Echinodermata The Echinodermata play a minor role in calcium carbonate production on the central west Florida shelf. These familiar, exclusively marine, benthic animals are widely distributed at all depths, latitudes and temperatures in the Gulf of Mexico as well as other oceans and are among the most abundant sea floor animals (MAFLA 1974, 1975-1976; Blake et al., 1995; Brusca and Brusca, 1990) The generally epifaunal, ubiquitous echinoderms display remarkable continuity on the west Florida shelf (MAFLA, 1974). They are intolerant of low salinity and are generally photonegative (Macginitie and Macginitie, 1968; Blake et al., 1995). The echinoderms are distinguished by their radially symmetric, calcitic endoskeleton. This endoskeleton is composed of separate plates of calcium carbonate, each plate a single crystal of calcite (Binyon, 1972; Brusca and Brusca, 1990). The fossil record, both in sedimentary deposits and rock formations, indicates over 13,000 species ofwell-preserved calcitic echinoderm skeletons (Vinogradov 1953; Raup, 1966). The five classes of echinoderms, Asteroidea, Crinoidea, Echinoidea, Holothuroidea and Ophiuroidea, contain approximately 7,000 living species, more than 74 of which are represented in the study region (Vinogradov, 1953; MAFLA 1974, 1975-41

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1976; Brusca and Brusca, 1990; Blake et al., 1995). These species include the feather stars, basket stars, brittle stars, sea stars, sea urchins, sand dollars and sea cucumbers (MAFLA 1975-1976; Brusca and Brusca, 1990). They may range in size from less than one centimeter to more than one meter in diameter (Vinogradov, 1953; Brusca and Brusca, 1990). Echinoderms exhibit different feeding behaviors that vary by species, food availability and bottom composition (Ebert, 1968). The carnivorous asteroids continuously feed as opportunistic predators or scavengers primarily on calcareous organisms and are abundant along the gulf coast of Florida (Giese, 1966; Macginitie and Macginitie, 1968, Dehn, 1980; Brusca and Brusca, 1990). The less dominant crinoids are suspension feeders (Brusca and Brusca, 1990). The holothuroids, although capable of selective deposit feeding, often feed by filter suspension and can be restricted to low-energy environments (Lawrence and Kafri, 1979; Brusca and Brusca, 1990). Both the crinoids and holothuroids tend to be inactive, almost sessile, and may have a great effect on the sea floor by feeding upon the organic content of the substrate, plankton and detritus (Macginitie and Macginitie, 1968). The highly competitive, well-adapted echinoids, although primarily herbivores, maximize their feeding strategy by being potential suspension feeders, deposit feeders, detritivores, predators or facultative omnivores (Giese, 1966; Lawrence and Kafri, 1979; Brusca and Brusca, 1990). The active ophiuroids feed by predation, selective deposit feedings, as detritivores and by suspension feeding (Macginitie and Macginitie, 1968; Brusca and Brusca, 1990). This variability among all the echinoderms allows adaptation by many different species in different environments with variable food supplies. 42

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The echinoderms of the west Florida shelf are highly patchy in distribution and biomass, with biomass ranging from 0 g m-2 to 230 g m-2 among the stations sampled (MAFLA, 1974, 1975-1976; Blake et al., 1995). This large range in distribution and biomass is attributed to a combination of factors characteristic of such patchiness. These factors include food supply and availability; feeding type; defensive tactics; substrate type; migration; spawning; mobility; weather; competition for space, for food and with other animals; size of animals; predation; aggregating behavior; and type and limitations of sampling methods (Ebert, 1968; Lane, 1977; Lawrence, 1978). Most echinoderms are gregarious in nature and many aggregations are composed of multiple species of echinoderms (Reese, 1966). These adaptations and strategies among the echinoderms likely optimize their survival, growth and reproduction rates, yet they limit the ability to quantify the biomass of the echinoderms of the study region by the restriction of the box core sample technique. The following production estimations should be considered a range of the total contribution to sediments by the echinoderms in the region. Methods Categorization of Organisms The five classes of echinoderms are all represented in the west Florida shelf biomass yet with unequal distribution. None of the data sets provide a breakdown by species or class of the macroinfauna sampled. Dominant taxa have been noted for the 1974 and 1975-1976 series ofMAFLA cruises and the MMS cruises. They are found in Appendix 27. 43

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Data Used Biomass values from the macrofauna! samples are available for both transects from the 1974 MAFLA cruises, two transects from the summer 1975 MAFLA cruise and from the MMS cruises of 1992-1994. All samples were obtained from 21.3 em x 30.5 em box cores with 32 em of maximum penetration. The box core penetration is limited by the composition of the bottom and substrate and may vary between stations (MAFLA, 1974). As for the molluscs, a faunal list was prepared from the screened box core samples for the dominant macroinfauna. Nine replicate box cores taken and analyzed per station represent 0.48 m2 of sea floor of the 1974 cruises, 0.54 m2 of sea floor for the 1975-1976 MAFLA cruises, and 0.57 m2 from the MMS cruises of 1992-1994. Echinoderm biomass figures are expressed in grams wet weight per square meter of sea floor. Calculations To estimate the calcium carbonate contribution of the echinoderms, many factors must be addressed. Growth rate information is not useful as it varies by area, season, temperature, food supply, reproductive cycle, and size of the organism (Ebert, 1968; Halpern, 1970; Crump and Emson, 1978; Dehn, 1980). Growth rates also are impacted by regeneration in damaged animals and negative growth periods (Ebert, 1968; Binyon, 1972; Dehn, 1980). Annual carbonate production is evaluated by estimating the amount of calcium carbonate in the echinoderm standing crop expressed as a percent of the wet weight biomass multiplied by the turnover rate or mortality. 44

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Chemical Composition Of the 7,000 extant echinoderm species, less than 10% have been chemically analyzed (Vinogradov, 1953; Raup, 1966). Chemical composition maintains a certain unity for the echinoderms with all the Echinodermata possessing a magnesium-calcium skeleton that comprises the majority of the organism's dry weight (Vinogradov, 1953). For calcium carbonate production rates, the amount ofCaC03 content ofthe organism as a percent of the biomass must be estimated. Chemical composition is a function of many factors including species, position in the skeleton, sea water temperature, season, size of individual and reproductive cycle (Vinogradov, 1953; Raup 1966; Binyon, 1972). Control ofresource allocation allows better adaptation for each organism during periods of somatic growth, reproductive growth, and regeneration with components varying considerably in size and composition during these life cycles (Giese, 1966; Lawrence and Ellwood, 1991). Therefore, no true correlation exists between body size or age or geography, and the organic composition of the body wall and carbonate composition are not constant within the animal (Binyon, 1972; Sibuet and Lawrence 1981 ; Lawrence and Guille, 1982) Carbonate dominates th e skeletal material, the majority is calcium carbonate and 5% to 15% is magnesium carbonate (Vinogradov, 1953; Binyon, 1972). The average echinoderm contains between 65% and 75% water with the holothuroids containing much more (Vinogradov, 1953; Binyon, 1972) The w a t e r content within the body does vary greatly between components with an average of 40% water content of the body wall (Giese, 1966) 45

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The body wall varies between 30% and 88% of the total wet weight for most species analyzed. Of the dry weight, echinoids contain the higher percentage of ash, up to 89%, the ophiuroids around 70%, asteroids 67% and holothuroids 3 7% ash for the species analyzed (Vinogradov, 1953; Giese, 1966; Binyon, 1972; Moore, 1972; Ebert, 1973, 1975; Lane, 1977; Lawrence and Guille, 1982; Lawrence and Bazhin, 1998). Again, these values are merely averages of available data for a multiple number of species, some identified within the MAFLA area and some that were not. The amount ofCaC03 in the ash is derived from Vinogradov's (1953) extensive compilation of chemical composition for a large number of species. An estimate of CaC03 as percent of the total wet weight biomass was calculated as the product of percent dry weight of the organism and the percent ash of that dry weight multiplied by the percent CaC03 in the ash. These values were then averaged by species identified in the sampled areas For the dominant species, 41% were asteroids, 38% echinoids, and 21% ophiuroids. A lack of consistency in the presentation of the available chemical analyses and the lack of chemical analyses on many species complicates comparisons Some analyses present body wall weight in p e rcent dry weight, some in ash residu e If the water content is not also provided it does not allow for a reliable method to evaluate the carbonate content. The estimations made here and summarized in Table 7 therefore, are broad averages incorporating a wide range of composition values for many org anisms in different environments. 46

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Table 7 Echinodenn chemical composition t':S t':S t':S Q) Q) Q) t':S "'t:: "'t:: "'t:: ...... '8 ...... ...... 0 0 0 E-< Q) s:: !:l to-. Sources ...... :.a :.a Cll Q) < (,) 8" ..!:: Cl) Water Content 66% 62% 63% Dry Weight 34% Vinogradov 1953 38% 37% Giese 1966 Ash in % Dry Weight 67% 85% 72% Binyon 1972 Moore 1972 % CaC03 in Ash 86% 88% 88% Ebert 1973, 1975 CaC03 As % Biomass 20% 28% 23% Lane 1977 Lawrence & Guille 1982 % ofWest Florida Shelf Species 41% 38% 21% Lawrence & Bazhin 1998 % ofProduction Rate 8% 11% 5% 24% Turnover Rates Turnover rate is a measure of the estimated life span until mortality. In a dynamic biological community, it is not easy to quantify either age or longevity for the echinoderms (Ebert, 1973 ). Size is a poor indicator of age and it is difficult to separate age classes (Binyon, 1972; Ebert, 1973; Dehn 1980). Ages for echinoderms have been recorded greater than 100 years (Lawrence and Bazhin, 1998). Turnover rate averages have been noted from a rate of0.83 estimating survival at less than one year to a rate of 0 1 indicating average length of life at 10 years (Smith, 47

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1971; Binyon, 1972, Chave et al., 1972; Ebert, 1973; Ebert, 1975; Lawrence and Bazhin, 1998). An average for the west Florida shelf region is 0.26 yr-1 approximately four years. Echinoderm Calculations Wet weight biomass values from the three different series of cruises were multiplied by the percent of calcium carbonate of the biomass and then by the turnover rate. Thus, carbonate production rates were estimated as: Echinoderm Biomass Wet Weighting m-2 x 24% CaC03 x .26 yr-1 =Calcium Carbonate Production Rate in g CaC03 m-2 yr-1 The rates are summarized in Table 8. Table 8 Echinoderm production rates for the central west Florida shelf R ange Average 197 4 Transect II 0.01-2.2 0.4 197 4 Transect I 0-14.4 1.6 197 4 Average 0-14.4 1.0 1975-1976 Transect I 0-9.1 1.9 197 4-197 6 Transect II 0-3.2 0.6 1975-1976 Average 0-9.1 1.2 MMS Site I 0-4.2 1.0 MMS Site II 0.01-6.5 1.3 MMS Site III 0.050.15 0.1 MMS Site IV 0.2-13 6.6 MMS Site Average 0-13 2.2 Florida Middle Ground 0.01-2.2 0.6 Average for shelf 014.4 1.3 48

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Discussion of Results The echinoderms, like the molluscs, show variability among the stations sampled. The total production estimation, 1.3 g CaC03 m-2 yr-1 is low compared with the other contributors, yet consistent across the shelf. The echinoderms are not a prominent constituent of the sediments (Brooks, 1981). Some trends have been noted with depth and a summary is found in Table 9. Table 9 Echinoderm production summary by depth range Depth Range 6-20m 29 -54m 90 -189m Florida Middle Ground Total for Shelf Shelf Average 3.01 0.83 0.02 0.62 1.3 The estimated production rates decrease with increasing depth. Production by echinoderms appears to be less for the Florida Middle Ground than the surrounding shelf. Observations and data have shown an increase in food availability on the Florida Middle Ground, which should affect an increase in echinoderm abundance. The biomass numbers for the echinoderms may be underreported for the region again owing in part to the limitations of the box core sampling method, the nature of their patchy distribution, and cryptic species hiding in holes in the hard substrate. 49

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CHAPTER VI CORAL AND CALCAREOUS ALGAL PRODUCTION Coral and calcareous algae are known to be prominent carbonate producers in reef environments (Chave et al., 1972; Land, 1979; Bosence, 1989; Hubbard et al., 1990; Milliman, 1993). On the central west Florida shelf, they also contribute to the carbonate produced, yet not in large reef-building communities. The species identified are of West Indian-Caribbean affinities (Cheney and Dyer, 1974; MAFLA, 1974). The factors influencing distribution, growth and calcification rates are intricately connected for these flora and fauna. Algae Calcium carbonate secreting algae are widely distributed in all seas, including this study area in the eastern GulfofMexico (Vinogradov, 1953). The calcareous algae found in warm tropical seas concentrate calcium carbonate quite intensively (Vinogradov, 1953). The benthic calcareous algae produce significant amounts of carbonate in marine systems in both recent and ancient times (Stockman et al., 1967; Chave et al., 1972; Neumann and Land, 1975; Milliman, 1993; Freile et al., 1995). In addition to the production of substantial amounts of carbonate and sediment, calcareous algae also help to shape the benthic environment as skeletal framework builders and by integration of organism and sediment through trapping, binding, baffling, cementing and encrusting ofthe substrate (Ginsburg et al., 1971; Milliman, 1993; Freile et al., 1995). 50

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A diverse assemblage of benthic algae with tropical and subtropical affinities occurs at all euphotic depths in the eastern GulfofMexico (Cheney and Dyer, 1974). The dominant carbonate secreting classes of algae on the central west Florida shelf are the Chlorophyta, Phaeophyta and Rhodophyta (MAFLA, 1974; MAFLA 1975-1976). The algae identified in the study region are summarized in Appendix 30 The Chlorophytes are distinguished by their green color, fast growth rates and their prolific production (Milliman, 1974; Freile et al., 1995). They thrive in temperate and subtropical environments (Milliman, 1974). Chlorophytes are usually most dense at depths less than 45 m but the calcareous species may extend to depths exceeding 400 m (Vinogradov, 1953; Ginsburg et al. 1971; Cheney and Dyer, 1974; Milliman, 1974; Freile et al., 1995). Significant beds of sea grasses and algae have been observed off the transects in the 1974 cruises (MAFLA, 1974). Halimeda, one ofthe most highly calcified of the Chlorophytes, can be found in very dense patches or meadows in loose sediment and hard bottoms producing much carbonate material as well as providing substrate for a substantive epibiont community (Milliman, 1974; Neumann and Land, 1975; Freile et al., 1995). The Rhodophyta or red algae are the most cosmopolitan of the calcareous benthic algae found commonly in tropical waters and at greater depths and latitudes (Ginsburg et al., 1971; Milliman, 1974). Known for their rigid skeletons, they primarily act as encrusters, cementers and prominent sediment producers in the benthic community (Milliman, 1974). The encrusting coralline algal species dominate at increasing depths and are more readily found on hard substrates (Milliman, 1974). 51

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The brown algae, Phaeophyta, although common, do not make a major carbonate contribution; Pad ina is the most significant of the calcareous species on the central shelf (Vinogradov, 1953; Milliman, 1974). Also listed are the Cyanophyceae or blue-green algae (now known as cyanobacteria) that have been identified. The cyanobacteria may contribute to carbonate production but with no information available no assessment will be made (Ginsburg et al., 1971; Neumann and Land, 1975; Yates and Robbins, 1998). Growth rate for the benthic algal species is influenced and controlled by a variety of environmental factors (Kleypas, 1997). The parameters most linked with growth rate include temperature, light intensity and quality, freshwater input, water circulation, nutrient availability, season and predation pressure (Stockman et al., 1967; Ginsburg et al., 1971; Payri, 1995). Increase in nutrient availability provides an advantage for algal growth over coral growth, yet the increase in nutrients in the water column may diminish the available light and thus negatively affect the production (Cheney and Dyer, 1974; Hallock, 1988; Canals and Ballesteros, 1997; Kleypas, 1997). Growth rate trends show an inverse relationship with depth (Hubbard et al., 1990). The growth rate for most algae is not well known and quantification is further complicated by the irregular growth spurts that have been documented (Colinvaux et al., 1965; Neumann and Land, 1975). Distribution of the calcareous algae depends upon many abiotic and biotic factors. Temperature; light intensity, quality and wavelength; competition and grazing pressure; and nutrient availability affect distribution (Ginsburg et al., 1971 ). Distribution is also linked with substrate, seasonal changes, water movement, salinity, carbon dioxide levels, and the effects of depth (Ginsburg et al., 1971; Cheney and Dyer, 1974; Payri, 1995; Canals and Ballesteros, 1997). 52

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Algal turnover rates show little uniformity. Ranges from a turnover of every 30 days to every six months have been documented (Stockman et al., 1967; Chave, et al., 1972; Neumann and Land, 1975). Some trends with depth have been shown as longevity may increase with increasing depth due to the decrease in growth rate and grazing pressure (Canals and Ballesteros, 1997). Grazing pressure can be more intense for the photophilic algae (Canals and Ballesteros, 1997). The assemblages vary across the shelf in response to variations in a combination of these factors (Canals and Ballesteros, 1997). Most of the calcareous algae have patchy abundance and vary temporally creating difficulties in quantifying and describing population densities and distribution (Stockman et al., 1967; Neumann and Land, 1975; Bosence, 1989; Yates and Robbins, 1998). Algal Productivity Of the algal assemblages, seagrass communities with coralline algal epibionts can produce among the largest amounts of carbonate in some nearshore, shallow habitats (Neumann and Land, 1975; Bosence, 1989; Frankovich and Zieman, 1994). Thalassia beds with their algal and serpulid epibionts may contribute up to 500 g CaC03 m-2 yr-1 in Florida Bay (Bosence, 1989; Frankovich and Zieman, 1994). Halimeda meadows and bioherms can exceed even these values and produce upwards of 3,000 g CaC03 m-2 yr -1 at greater depths, 20 to 100m (Milliman, 1993; Freile et al., 1995). These environments are not typical of the central west Florida shelf and no estimate will be made for these types of communities. From comparison of production rates, some trends have been demonstrated. Change in light intensity controls photosynthesis and consequently productivity, and, as 53

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light intensity is attenuated with increasing depth, it results in a decrease in productivity (Hubbard et al., 1990; Murray, 1991; Canals and Ballesteros, 1997). Although not a direct linear function of depth, much of the growth rate and productivity is light controlled and limited, and calcification rates can be as much as three times higher in higher light conditions (Hubbard et el., 1990; Murray, 1991; Payri, 1995; Canals and Ballesteros, 1997; Kleypas, 1997). Calcification rates are not always directly linked with photosynthesis as some algae have adapted at depths to lower levels oflight (Ginsburg et al., 1971; Milliman, 197 4; Canals and Ballesteros, 1997). The abrupt drop in carbonate production with increasing depth is largely attributed to the reduction in light (Hubbard et al., 1990). Productivity and longevity may increase at greater depth for some species due to a reduction in grazing pressure upon the photophilic algae (Land, 1979; Hubbard et al., 1990; Canals and Ballesteros, 1997). Algal production changes noticeably with changes in substrate (Milliman, 1974; Canals and Ballesteros, 1997). Rocky bottom areas enjoy greater amounts of algal carbonate productivity than the sedimented areas, notably among the encrusting species (Milliman, 1974; Canals and Ballesteros, 1997). High carbonate content sediments have an increase in productivity over the finer, sandier materials (Canals and Ballesteros, 1997). Corals Over three dozen Octocorallian and Scleractinian corals have been identified from the MAFLA cruises of 1974 and 1975-1976. These octocorals and stony corals found in 54

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Appendix 31 were archived and listed by transect only, with a large number found in Transect II of 1974 and Transect III of 1975-1976, the Florida Middle Ground area. The colonial Octocorallia include the sea fans, sea whips and sea rods and their polyps are comprised of eight calcareous sclerite tentacles. The also colonial Scleractinians, with abundant symbiotic zooxanthellae, have delicate to massive calcareous exoskeletons capable of tremendous reef formations (Brusca and Brusca, 1990; Allaby, 1992; Humann, 1994). Coral Productivity Within the central shelf region, corals are not evenly distributed. They are common in the Florida Middle Ground and very rare and patchy in the other shelf areas (MAFLA, 1974, 1975-1976). The eggs and larvae of these tropical and sub-tropical coral species likely are transported on to the shelfby the loop waters (MAFLA, 1974). Light, effects of depth, turbidity, sedimentation rate, temperature, water motion, seasonal effects, substrate and geography, all control growth rates and distribution (Vaughan, 1911, 1915, 1917; Goreau, 1959; Shinn, 1966; Smith and Kinsey, 1976; Smith, 1978; Land, 1979; Gladfelter, 1984; Hubbard and Scaturo, 1985; Hubbard et al., 1990; Heiss, 1995). One benefit to coral growth on the shelf is the lack of sediment load (Vaughan, 1917; Hallock, 1997) As depth increases, growth rate and calcification rates both decrease for coral. Most coral reef formation occurs in depths ofO to 30m, some coral grow in the 30 to 74 m range and a few species grow at greater than 75 m depth but at a dramatically lowered rate and occurrence (Vaughan, 1917; Shinn, 1966; Smith and Kinsey, 1976; Smith, 1978; Hubbard and Scaturo, 1985; Heiss, 1995). 55

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Turbidity and water motion play roles in coral growth. Production rates decline with a decrease in water energy. Storm events and currents change light, nutrient availability, and sedimentation rate and may effect temperature and salinity (Vaughan, 1911, 1915; Smith and Kinsey, 1976; Hubbard and Scaturo, 1985). Vaughan (1911, 1915, 1917) noted comparatively lower growth rates in Gulf of Mexico West Indian species of coral than in the Pacific assemblages, as well as an increase in coral growth on firm or rocky substrates. Temperature controls much of the coral growth and calcification with an ideal range of 26C to 28C, average minimum of 18C, and death at a sudden drop below 13C (Vaughan, 1911; Shinn, 1966; Gladfelter, 1984). The bottom temperatures in this central west Florida shelf region average 17C to 29C in summer, 17C to 27C in fall, and 12C to 21C in the winter (MAFLA, 1975-1976). Seasonal changes affect most of the growth rate parameters like temperature, water motion, nutrient availability and change in light intensity and length of light exposure. Most agree that light is the key control in coral distribution and growth and an increase in depth exponentially decreases coral productivity (Vaughan, 1911; Goreau, 1959; Land, 1979; Gladfelter, 1984; Hubbard and Scaturo, 1985; Hubbard et al., 1990). Chemical Composition Chemical and water content of the calcareous algae and coral vary by species and class, with the season of the year, age ofthe organism and vertical distribution (Vinogradov, 1953). From the available data, water content ranges from 58% to 85% for most ofthe benthic algae (Vinogradov, 1953). Some species' calcium carbonate content averages 25% ofthe wet weight of the plants (Bosence, 1989). For the coralline red 56

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algae, ash residue may exceed 50% of the wet weight with upwards of 60% to 90% of the ash being calcium carbonate (Vinogradov, 1953). The calcium carbonate content of the chlorophytes ranges from 45% to 65% of the dry weight (Neumann and Land, 1975). Corals and Halimeda, one of the more prominent algal species in the study region, may contain as much as 99% calcium carbonate ofthe ash residue (Vaughan, 1911, 1917; Vinogradov, 1953; Goreau, 1959). Carbonate Productivity Coral and calcareous algae are known to be major carbonate producers in reef and other marine environments and can, in some habitats, be locally very important to the total carbonate produced with production rates approaching 104 g CaC03 m-2 yr-1 (Chave et al., 1972; Neumann and Land, 1975; Land, 1979; Bosence, 1989; Hubbard et al., 1990; Milliman, 1993; Freile et al., 1995; Hallock, 1997; Yates and Robbins, 1998). Reefs play a key role in the global carbonate budget and have been extensively researched (Le Carnpion-Alsumard et al., 1993; Milliman, 1993; Kleypas, 1997). Modem reefs precipitate enough calcium carbonate to keep up with the rise in sea level (Chave et al., 1972; Smith and Kinsey, 1976). Few quantifications of coral and algal carbonate productivity for shelf systems have been made, estimating rates has proved very difficult (Stockman et al., 1967; Chave et al. 1972; Land, 1979; Bosence, 1989; Hubbard et al., 1990; Yates and Robbins, 1998). No biomass quantifications or descriptions by station were present e d for the MAFLA or MMS cruises. Lacking any data for this region, the calcium carbonate production rate must be adapted from rates calculated by other studies summarized in Tables 10 and 11. 57

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Table 10 Summary of algal production rates Assemblage Production Depth Location Source g CaC03 m 2 yr-1 m Predominant Red and 100 <100 Mallorca-Menorca Canals and Green Algae shelf Ballesteros 1997 Northwestern Mediterranean Sea Coralline Algae 20 <40 Shelf-Edge Reef Hubbard et al. No Halimeda (1.65% of total System 1990 production ) U.S. Virgin Islands Halimeda 2400 20-40 Grand Bahama Bank Freile et al. 1995 100% Coverage Slope Benthic Algal Community 500 <50 Bank-Embayment Milliman 1993 Halimeda 3000 30-100 Bioherm Thalassia Epibionts 1.9 282.7 3 Florida Bay Frankovich and Coralline Red Algae Zieman 1994 dominant Thalassia Epibionts 81 Florida Bay Inner Bosence 1989 Penicillus 4 .83 Halimeda Trace Thalass i a Epibionts 482 Florida Bay Open Penicillus 29.25 Halimeda 11 Red Coralline Algae 500-2500 Hawaii Littler 1971 0 7000 St. Croix A dey and Vassar 1974 Marine Lagoon 90-100 7 Abaco, Bahamas Neumann and Land 1975 Green Algae 100 Sand Reef Model Chave et al. 19 72 Red Algae 1000 Alg a l ridge Red Algal dominant 400 Subtidal California Borderland Smith 1971 Community P e n i cillus 3 .23 Florida Bay Stockman et al 25 Inner Reef Tract 1967 58

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For purposes of estimation, carbonate productivity is assumed to be inversely related to depth with the majority of production in less than 90 m; productivity increases on hard substrates and also carbonate-dominated sediments; and overall productivity will be higher in more oligotrophic waters. Table 11 Summary of coral production rates Assemblage Production Depth Location Source g CaC03 m"2 yr1 m Corals 1210 <40 Shelf-Edge Reef Hubbard et al. (93% of total Syst e m 1990 production) U.S. Virgin Islands Coral Reefs 1500 Milliman 1993 Porites 513.2 Florida Bay Bosence 1989 Fore-Reef Slope 600 20-60 N. Jamaican Island Land 1979 (33% of total Slope production) Shallow, sand flat with 410 Sand Reef Model Chave et al. 1972 calcareous algae Coral Reefs 800 5-6 Protected Smith and Environments Kinsey 1976 The central west Florida shelf varies in depth and substrate. Change occurs across the shelf from low and high relief rocks to sediments high in carbonate material to hard compacted sand to soft silt shell rubble or fine sand (MAFLA 1974). In an attempt to address as many ofthe variables as possible, the shelf region is divided into four different depth and sediment composition zones: 6 to 29 m depths; 30 to 54 m d e pths; 90 m and grea ter depths; and th e F lorida Middle Ground. The shallow es t zone with 14 stations, 12m average depth has the lowest average percent calcium 59

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carbonate in the sediment, 37%, and is sand-dominated. The sediments of the 22 stations of the 30 to 54 m zone average 40 m in depth and 94% CaC03 in the sediments and are a mixed community of carbonate sediments and Halimeda. The four stations greater than 90 m have sediments of94% CaC03 with an average depth of 156m. The nine stations of the Florida Middle Ground range from 34 to 44 m, 39m average, and have an average sediment carbonate content of 77%. A combination of rocky substrate, hard bottom and carbonate sediments makes up the Florida Middle Ground. No stations are between 55 and 89 m Sediment constituency can indicate benthic community composition (Land, 1979). The sediments of the Florida Middle Ground show 1.5% coralline algae skeletal fragments, a trace amount of Halimeda and 2.2% coral skeletal fragments, and, the surrounding shelf contains merely a trace of each (Brooks, 1981 ). The sediment analysis identified Halimeda fragments in each ofthe stations sampled for the 1974 MAFLA cruises, coralline algae in 66% of the stations sampled and coral fragments in 11% ofthe stations in this study region (MAFLA, 1974). Thus, algal productivity seems more consistent over the shelf area. The study area of the central west Florida shelf should have a lower rate of carbonate production than other coral and algal rich environments with similar assemblages for a variety of reasons, namely: a decreased amount and quality of light; lower average temperatures, greater depth; an increase in eutrophication; higher latitude; a lack of much firm and rocky substrate; a low energy environment; and patchy coverage. The rate of coral and algal production on the Florida Middle Ground is assumed to be higher than on the surrounding shelfbecause of the increased number and abundance of 60

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carbonate-producing coral and algal species observed, the increase in water clarity, the firm and rocky substrates observed, and the increase in algal and coral skeletal fragments found in the sediment (MAFLA, 1974, 1975-1976). A production rate of25% ofthe Florida Middle Ground for the shallow, sandy 6 to 29m zone and a rate of 10% for the 30 to 54 m zone is assumed. The coral species' carbonate production, sensitive to the combination of changes in light, temperature and an increase in nutrients, declines logarithmically with depth. The average rate of production by corals found in somewhat similar environments range from 102 to 10 3 g CaC03 m-2 yr-1 and average 850 g CaC03 m-2 yr-1 The Florida Middle Ground production, with an exponential reduction, should average 3 g CaC03 m-2 yr-1 For the 6 to 29m zone, a rate of0.75 g CaC03 m-2 yr-1 is estimated and 0.3 g CaC03 m-2 yr-1 for the 30 to 54 m zone. A zero value for the stations 90 m and greater will be assumed as the average annual production would be negligible. Algal productivity varies more linearly with depth. The average rate of production for several similar environments is approximately 250 g CaC03 m-2 yr-1 (see Table 1 0). The productivity for the Florida Middle Ground is estimated at 10% of that, 25 g CaC03 m-2 yr-1 ; 6.25 g CaC03 m-2 yr-1 for the 6 to 29 m zone; 2.5 g CaC03 m-2 yr-1 for the 30 to 54 m zone; and 0.25 g CaC03 m-2 yr-1 for the depths 90 m and greater. These results are found in Table 12. 61

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Table 12 Coral and algal production rates for the central west Florida shelf In g CaC03 m-2 yr-1 Zone Coral Algal Total Florida Middle Grounds 3 25 28 6 to 29m 0.75 6.25 7 30 to 54 m 0.3 2.5 2.8 0 0 0 62

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CHAPTER VII PRODUCTION BY OTHER ORGANISMS Several other types of calcium carbonate secreting organisms are worthy of mention. These locally may make a significant contribution yet do not contribute in a major way to the total shelf production rate (Milliman, 1993). Annelida Of the annelids, the polychaete class is extremely abundant across the central west Florida shelf(MAFLA, 1975-1976). The Family Serpulidae, tube worms, build hard, calcareous tubes into which the worm retreats (Brusca and Brusca, 1990; Humann, 1992; Aliani et al., 1995) The tubes make up an average of96% CaC03 ofthe ash residue (Vinogradov, 1953). These sedentary animals are suspension feeders (Brusca, and Brusca, 1990). Many of the serpulids are found on the shelf including the well-known Christmas tree worm, Spirobranchus giganteus, and are listed in Appendix 32 (MAFLA, 1974, 1975-1976; Humann, 1992; Blake et al., 1995). Polychaetes have been identified at every station of the MAFLA, MMS and the Project Hourglass 1965-1967 cruises. They rang e from 91 to 4,240 individuals per square meter and average over 1,000 individuals per square meter, with biomass ranging from 0.2 to 25 grams per square meter, averaging 5.5 grams per square meter wet weight 63

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(MAFLA, 1974, 1975-1976). The polychaetes average 14% of the total macroinfaunal biomass (MAFLA, 1974). Massive tube formations are possible as the worms may become quite densely packed (Vinogradov, 1953; Aliani et al., 1995). However, very few of the dominant species at any stations are of the serpulid family and these types of aggregations are very patchy and rare on the central west Florida shelf(MAFLA, 1974, 1975-1976, T. Perkins, personal communication). The carbonate production rate for the serpulids would merely be a trace across the shelf and no value will be included. Porifera Of the Porifera, Class Calcarea, the exclusively marine sponges, secrete a skeleton of calcium carbonate (de Laubenfels, 1953; Vinogradov, 1953; Bergquist, 1978). The Calcarea are usually limited to depths less than 100 m and to firm substrates (MAFLA, 1974; Bergquist, 1978). Water content, as expected for the sponges, is high, close to 90%; the ash residue, a mere 6% ofliving matter, is nearly 87% CaC03 (Hyman, 1940; Vinogradov, 1953). The Calcarea, like the Echinodermata, are high in MgC03 (Hyman, 1940; Vinogradov, 1953; Raup, 1966). The sponges play a major role in providing cover and concealment of other organisms as well as playing host to bacteria and cyanophytes (MAFLA, 1974; Bergquist, 1978; Brusca and Brusca, 1990). Sponges can effectively excavate and break down calcareous material that is expelled in their exhalent stream (Bergquist, 1978). Of the nearly 200 species of sponge located on the central west Florida shelf, only eight Calcispongiae species were identified, most on the Florida Middle Ground where a higher sponge density and diversity occurs (MAFLA, 1974). Yet sponge spicules have 64

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been recorded in the sediment analysis at every site (MAFLA, 1974). Again, none are dominant on the shelf and they probably contribute only a trace amount of carbonate to the system. Bryozoans The colonial bryozoans or ectoprocts found on the shelf also are calcified (Brusca and Brusca, 1990; Humann, 1992). Skeletal portions ofbryozoans are found in the sediments of the shelf in trace amounts. The contributions by the bryozoans are usually disregarded (Neumann and Land, 1975; Bosence, 1989). Crustacea Benthic Crustaceans The ubiquitous crustaceans are found at all stations on the central west Florida shelf from benthic community to the water column. These communities' composition changes across the shelf (Hopkins et al., 1981 ). The most highly calcified of the crustaceans are the Cirripedia, whose shells are chiefly CaC03 97% (Vinogradov, 1953; Brusca and Brusca, 1990). These animals are the sole significant sediment constituent found only on the Florida Middle Ground and not the surrounding shelf(Brooks, 1981). Sediment analysis reveals over 8% barnacle skeletal fragments on the Florida Middle Ground and a trace amount elsewhere (Brooks, 1981 ). The sessile barnacles may produce measurable amounts of carbonate to the system and should not be ignored. Lacking any data other than sediment constituency, a gross estimation for the Florida Middle Ground stations is 25% of the production of the molluscs, which comprise 36% of the skeletal fragments in the sediment in the region. 65

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Thus an additional 0.5 g CaC03 m-2 yr-1 is added to the nine stations of the Florida Middle Ground. Water Column Biomass The MAFLA cruise identified 104 different zooplankton species of copepods, cladocerans, amphipods, ostracods, mysids, isopods, decapods, euphausids, and thalliaceans (MAFLA, 1975-1976). Zooplankton biomass averaged 0.04 g m-3 dry matter over the cruise periods (MAFLA, 1974; MAFLA, 1975-1976) Many of the crustaceans have a predominantly chitinous, as opposed to calcitic, carapace (Vinogradov, 1953; Brusca and Brusca, 1990; Gunthorpe et al., 1990; Gore, 1992). For the carbonate secreting species, just over 15% of the dry matter is CaC03 (Vinogradov, 1953; Johnson and Hopkins, 1978) With no information regarding the species composition ofthe biomass, the distribution in the water column, the distribution across the shelf, the residence time in the region, or the turnover rate of the zooplankton, no attempt to quantify this contribution to the production rate should be made. Fish Fish otoliths are comprised of concentric layers of calcium carbonate and organic material, as are a small percentage ofthe bone and scales (Vinogradov, 1953; Pannella, 1980 ; Kingsmill, 1993). The approximate otolith weight to fish biomass weight ratio is 1 o-8 with 1 o-5 maximum (R. Wilson, personal communication). For the observed fish biomass of 6, 700 g wet wei ght and an annual turnover rate of 12% to 15%, the 1 o -3 to 1 o -6 g CaC03 m -2 yr-1 is a negligible value for the shelf production rate. The calcification rates vary with temperature, salinity, pH, and feeding activity (Pannella, 1980). 66

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Unicellular Production Microbial calcification may also make major contributions to the carbonate production rate, yet little is known ofthis contribution (Neumann and Land, 1975; Yates and Robbins, 1998). The fossil record indicates large-scale carbonate deposition by cyanobacteria and microalgae (Yates and Robbins, 1998). Whitings, a not totally explained phenomenon, are precipitations of CaC03 from algal blooms, coccolithoid algae; microbial calcification; chemical precipitations; or, unlikely, resuspension of carbonate sediment (Stockman et al., 1967; Neumann and Land, 1975; MacGinitie and MacGinitie, 1968; Takano et al., 1994; Verrecchia et al., 1995; Yates and Robbins, 1998). These dramatic events may spike the carbonate production rate but for a very limited area. At this point, not enough data exist to quantify effects on this shelf. Anthropogenic influences like an increase in bicarbonate from wastewater discharge also will allow precipitation of CaC03 (Heyl, 1992). Bacteria also may enhance the deposition ofCaC03 (Dexter and Lin, 1991). Again, worthy of mention, yet no tangible values are available to include in the production estimations. 67

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CHAPTER VIII FLORIDA MIDDLE GROUND PRODUCTION The Florida Middle Ground is the northernmost hermatypic coral community in the eastern GulfofMexico (MAFLA, 1975-1976; Brooks, 1981; GMFMC, 1982). This reef community is recognized as an area of unique biological sensitivity (MAFLA, 19751976; Brooks, 1981 ). The nearly 700 km2 located 160 km north-northwest of Tampa, Florida, consists of discontinuous limestone outcroppings with relief of 2 to 10 m over depths of 25 to 65 m (Cheney and Dyer, 1974; Brooks, 1981 ). Two major parallel ridges are evident with broad valleys between them (Brooks, 1981 ). The Florida Middle Ground lies in the midst of a complex mixing area between the three significant water masses, the Loop Current, the West Florida Estuarine Gyre, and the Florida Bay Waters, with upwelling common in the area (Cheney and Dyer, 1974; MAFLA, 1974; Brooks, 1981). The Florida Middle Ground enjoys steady conditions of clear water, stable salinity and light penetration, and a temperature range of 16 to 26C with some thermal stratification (Cheney and Dyer, 1974; Brooks, 1981 ). The flora and fauna on the Florida Middle Ground are displaced far north of their usual limits. The outcroppings are covered with shell rock and living and dead corals of the north tropical, Caribbean affinities (Cheney and Dyer 1974). The diverse assemblages are rich in algal species, and hard and soft coral occur in patchy abundances with few species found on the loose sand or shell (MAFLA, 1974). 68

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The carbonate content of the sediments is typical of other continental shelves and the sediment constituents resemble those of the central west Florida shelf, not a typical coral reef environment (Brooks, 1981 ). Carbonate content decreases away from the Florida Middle Ground indicating a decline in carbonate production, with no significant relationship with depth noted (Brooks, 1981 ). Molluscan fragments dominate and the notable difference between the Florida Middle Ground and the surrounding shelf is the presence of Cirripedia due to the abundance of hard substrate for attachment (Brooks, 1981 ). Production for the Florida Middle Ground is summarized in Table 13, totals by station are found in Appendix 33. The range and average production for the nine stations of the Florida Middle Ground are included with the production for six stations immediately surrounding the Florida Middle Ground, as well as the average for all the stations of the central west Florida shelf except those of the Florida Middle Ground, and the entire shelf. The two distinct differences between the production on the Florida Middle Ground and the rest of the shelf are the increase in molluscan, algal, coral and barnacle production, and the decrease in the production for the foraminifera. The Florida Middle Ground's carbonate-producing over the surrounding area is the hard, elevated substrate and increase in water motion (Brooks, 1981 ). This allows favorable conditions for the tropical coral and algal species likely recruited from the Loop Current yet no active reef growth has been observed and coral and algal contribution is minimal (See Table 14) (Brooks, 1981). 69

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An anticipated dramatic change in calcium carbonate production rates for the Florida Middle Ground is not observed. Production is higher but similar for the Florida Middle Ground and the central west Florida shelf. These production values may be largely underestimated partially due to the limitations of the box core. The rugged bathymetry likely includes more surface area upon which calcareous organisms may live, increasing production rates but hampering the ability to quantify those rates. The overhangs, crevices and holes should host a variety of organisms that may be inadequately represented here (MAFLA, 1974). The patchy distribution of the organisms may also affect the biomass values. Table 13 Carbonate production rates for the Florida Middle Ground Florida Middle Surrounding Shelf without Central West Ground Shelf Florida Middle Florida Shelf Ground Range Avg Range Avg Range Avg Range Avg Foraminifera 9-53 24 32-86 52 6-124 41 6-124 38 Micromollusca 19-84 53 20-73 42 0-130 37 0-130 40 Macro mollusca 0.2-26 8 0.6-9 3 0 1-391 18 0.1-391 16 Mollusca 27-98 61 23-74 45 0.5-408 55 0.5-408 56 Echinodermata 0.01-2 1 0-0.83 0.3 0-14 1 0-14 1 Algae 25-25 25 3-3 3 0.3-6 4 0.3-25 8 Coral 3-3 3 0-0.3 0.3 0-1 0.4 0-3 1 Cirripedia 0.5 0.5 0 0 0 0 0-0.5 0.1 Total 82-165 114 73-123 100 25-434 101 25-434 103 Depth in m 34-44 39 37-54 46 6-189 42 6-189 42 Carb. In Sed. 10-98 77 78-92 86 10-98 71 10-98 72 70

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The sediments, summarized in Table 14, more closely resemble a carbonate continental shelf than a coral reef (Brooks, 1981 ) Production and deposits in the sediment are very similar to those on the shelf, molluscs dominated with significant amounts ofbamacle fragments (Brooks, 1981) The Florida Middle Ground has every indication of a transitional environment. This fossil reef system may be approaching assimilation into the central west Florida shelf (Brooks, 1981 ). Table 14 Sediment constituents modified from Brooks ( 1981) Skeletal Fragments Florida Middle Ground Surrounding West Florida Shelf Molluscs 30% 47% Foraminifera 3% 10% Coral 2% Trace Coralline Algae 1% Trace Halimeda Trace Trace Cirripedia 8% Trace Misc. Skeletal 15% The Florida Middle Ground is known to be an exceptional and productive fishing ground for commercial and sport fishing, as well as a dive destination (Cheney and Dyer, 1974 ; MAFLA, 1974; Brooks, 1981) Potentially small changes in the environment could create significant impact upon this biological system. However, carbonate production may not be severely impacted as production at 114 g CaC03 m-2 yr -1 is estimated to be around 10% greater for the Florida Middle Ground than for the other sites on the central west Florida sh e l f calculated at 103 g C a C03 m -2 yr-1 The stations 71

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immediately surrounding the Middle Ground enjoy nearly the same productivity as the reef system itself, 100 g CaC03 m-2 yr-1 albeit by a different assemblage of organisms. 72

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CHAPTER IX RESULTS OF CARBONATE PRODUCTION RATES FOR THE CENTRAL WEST FLORIDA SHELF Carbonate production rates have been estimated for some of the benthic organisms found on the central west Florida shelf. The results for all 48 stations sampled on the shelf are summarized in Table 15. For stations that were lacking data, an estimate is included from averages of other stations with similar depth and substrate. A complete summary by station is found in Appendix 34. Table 15 Calcium carbonate production rates CaC03 Production on the Central West Florida Shelf Range Average In g CaC03 m-2 yr-1 % of Production % of Sediments Foraminifera 6-124 38 37% 10% Micromollusca 0-130 40 39% Macro mollusca 0.1 -391 16 15% Mollusca 0.5-408 56 54% 47% Echinodermata 0 14 1 0.1% Trace Algae 0.25-25 8 8% Trace Coral 0 3 1 0.1% Trace Cirripedia 0-0.5 0.1 0.01% Trace Total 25434 103 Depth in m 6-189 42 Carbonate in Sediments 10-98 72 73

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CHAPTER X DISCUSSION OF RESULTS The average carbonate production rate is estimated at 103 g CaC03 m-2 yr-1 for the central west Florida continental shelf. This represents estimations obtained from 48 stations of diverse environments. The rates range in value from 25 to 434 g CaC03 m -2 yr-1 Carbonate production rates vary between stations, by season, by year and with each different assemblage of carbonate-secreting organisms. Regrettably, few definite trends are noted. The rates are graphically depicted in Figure 7 and no predictable patterns are delineated by depth, sediment-facies change or latitude. Several biotopes were identified within the study region including areas of hard compacted sand with silt, shell rubble, high-relief and low-relief rock ridges, coarse and hard packed sand, and soft and silty sand (MAFLA, 1974). Complex biotope changes occur over short distances and each zone intergrades into neighboring zones lacking any sharply defined boundaries (MAFLA, 1974; Lyons and Collard, 1974). Each change in biotope brings a consequent change in substrate, the infaunal benthic community composition, species distribution and species density. The communities across the shelf respond to a multitude of abiotic, biotic and chemical factors including: light intensity, quality and wavelength; substrate type; temperature; water clarity and movement; salinity; carbon dioxide ; available nutrients; 74

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Figure 7 \;.::11:.-i r u <11 ... iu .... r. ........ I I l'-"lllldard TIM" :t,r .\ \ (ll lc,i._ I L I II -\Stations with Calcium Carbonate Production Rates 0 l ) .. rti. : ,.. ..... J. 75 g Total Production ... 25 35 ... 36-52 ... 53 -61 ... 62-84 8597 98 110 111 133 .+ 166 .... ... 167 196 .... t ... 197 434

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season; year; depth; species type; competition and predation; disease; and age of organism (Ginsburg et al. 1971; Lawrence, 1978; Bishof, 1980) Different areas precipitate calcium carbonate at different rates (Langer et al., 1997). With a suite of bathymetric, geographic and seasonal variables influencing community structure, species abundance, diversity and density varied greatly between stations (MAFLA, 1974; MAFLA, 1975-1976). Assemblage composition, standing crop, turnover rates and growth rates are largely habitat-type dependent (Bishof, 1980; Ivany et al., 1994) Difficulties arise in even attempting to assign a species to a specific habitat (Buzas et al. 1993) Yet, a healthy, diverse infaunal benthic community is composed of just such a spectrum of species that do vary greatly in habitat, modes of feeding and tolerance of environmental stresses (Blake et al., 1995). The highly patchy, mobile and relatively long-lived benthic organisms are considered good indicator organisms (Blake et al. 1995). Buzas et al. (2002) also observed significant differences among stations, years and seasons for foraminiferal densities in the Indian River Lagoon, Florida, and determined environm e ntal variables offered little illumination to th e v ari ability in densities. The observed variability of foraminiferal densities between stations was quite lar ge with changes by station, season and year with every possible variation occurring. They concluded that th e variations in d e nsities are as complicated as po ss ible. The Buzas et al. (2002) model suggests asynchronous or aperiodic pulsating patches that vary in space and time H owever no over all trend of density increase or d ecrease was obs e rv ed. L on gt e rm stability is thu s a chi e v e d throu gh short t e rm 76

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variability both spatially and temporally. They also suggest that foraminifera and most other marine organisms exhibit the same patterns and the concept of pulsating patches is relevant for benthic macrofauna} organisms as well (Buzas, 1995; Buzas et al., 2002). Therefore, although the data show differences between stations, this variability should be expected and indicative of a healthy, stable carbonate-producing system. The pulsating patches observed across the shelf suggest long-term stability and the carbonate production rate of 103 g CaC03 m-2 yr-1 may provide a reasonable long-term average for a first estimation. Carbonate-producing assemblages vary significantly between most stations in species composition and dominance as well as species densities yet the estimated carbonate production rates remain within an order of magnitude reflecting a remarkable uniformity. This also might indicate a cyclical continuity and stability across the shelf. Factors Affecting Evaluation of Estimations An evaluation of the first order estimated production rates presented must address the many sources of error possible for such analysis. Sampling Technique The first consideration that arises is in the selection of stations Care was taken to choose a representative grouping of stations across several transects and the inclusion of the Florida Middle Ground. However, it is yet unknown if the 48 stations are indeed repr e sentativ e of the complete central west Florida shelf e nvironment. The box core technique introduces error in several ways. A bow wave is created that may wash away fauna from the edge of the core (Bishof, 1980). The box core is unabl e to pene t r ate hard substrat e s and depth o f penetration if d e p ende nt upon the typ e o f 77

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substrate (MAFLA, 1974; Bishof, 1980). The box core omits areas of great topography as it is unable to sample regions of relief, crevices, holes and overhangs, often areas of increased carbonate productivity. The data are greatly affected by the chance recovery of large, heavy specimens (MAFLA, 1974). The aggregation behavior of the large benthic organisms may lead to both underestimation if the patches are missed and overestimations should the box core collect an entire patch. The mobile benthic organisms may move into or out of the study area and may exhibit escape behavior from the box cores. Error may occur in the sorting of samples, live between dead; the processing of the samples; chemical analysis; depth of sample from the cores; classification and size sorting of the organisms; and in the identification of the organisms. Difficulties exist in the ability to collect live sample and the techniques used hampered recovery of some live specimens, especially of the micromollusca (Moore, 1972; MAFLA, 1974; MAFLA, 1975-1976). Organisms Omitted Due to a lack ofbiomass figures or densities available, several groups of carbonate-producing organisms are omitted from the production estimations. Planktonic production rates were not factored into the rates presented. Also, no value for production by the serpulid worms was calculated. Although very patchy in nature, their contribution could be locally important. The Porifera, sponges, are identified in the sediment analysis but not in the biomass figures. Therefore, evidence exists of a contribution of carbonate but no live specimen data from which to estimate it. 78

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Cyanobacteria may produce significant amounts of calcium carbonate yet remain an elusive subject (Neumann and Land, 1975) No estimation of the production or biomass value for the cyanophytes is available for these stations in the Gulf of Mexico. Therefore, as Neumann and Land (1975) have reported, if the contribution is minimized, it is only because of our ignorance of its importance. In addition to these organisms that have been intentionally neglected in the estimations, there may be some organisms inadvertently overlooked as they were not identified at the stations sampled. Unknown Factors To date, little data from the complex living benthic carbonate-producing shelf environments have been published, leaving much information yet unknown (MacGinitie and MacGinitie, 1968; Moore, 1972; Murray, 1991; Milliman, 1993; Blake et al., 1995). The unknowns encompass many areas especially the fields of biology, geology, chemistry, and climatology Certainly, an increased knowledge of the biology of the carbonate-secreting organisms would enhance the estimations for production. To better locate and identify all organisms present and to know the biomass, life history, recruitment strategies, growth and turnover rates and standing crops would improve production rates and pave the way for determining the sensitivity to how changes in their environment affect the ability to precipitate calcium carbonate. Geologically, the complete composition of the sediments, sedimentation and accumulation rates, and the age ofthe sediments have yet to be determined for the central west Florida shelf. Further analysis of the chemical composition of carbonate organisms 79

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also is required to improve production rate estimations Data of changes in the seawater chemistry would provide additional information as would the effects of carbonate dissolution at the sediment seawater interface. Missing, also, are the spatial and temporal effects of weather, storm events, and seasonal cycles on the benthos. Any efforts to fill in these blanks will doubtless advance the study of carbonate shelf production. Evaluation of Assumptions Made To complete an estimation for carbonate production across the shelf, many assumptions were made. The assumptions are somewhat data dependent as a different set of station data would have required different assumptions. The classification of the foraminifera by size mirrored the standards for the species presented. The micromollusc to macromollusc boundary is less clearly defined. Some have preferred to choose a 5 mm maximum for the micromolluscs rather than the 7 mm. The samples were impoverished with live material of any size so it may have not created a large source of error. Average chemical composition and growth rates are largely habitat dependent. Using archival data from many species located in many different environments at different latitudes likely only provides a range of composition and rates that might be found and was not specific to the Gulf of Mexico region. The grouping of species may also have been too broad. Although species composition and density within phylums were taken into consid era tion, the production rates should be more species-specific with better biomass analysis from the original samples. 80

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Comparison of the central west Florida shelf to other carbonate-rich systems was a best attempt to correlate this system with others. As more information of the subtleties of each environment is known, similar systems may be more easily recognized. Facing all of the numerous variables and the complexity of this marine environment, the same conclusion must be drawn as that by Stockman et al. (1967) when performing similar studies, namely, that it is impossible to evaluate quantitatively all the sources of error in the estimates of rates of production and accumulation. Thus stated, it also important to note that this estimation of production, 103 g CaC03 m2 yr-1 was created from actual data, observations and analyses made at nearly 50 stations on the central west Florida shelf. And, with no initial data or production rate estimates, there were no expectations about the predicted rate. This estimation provides a tool, a reference point and a working model from which to evaluate the shelf and to apply new data as it becomes available. Future Research The next steps in refining the carbonate production rate estimations should focus on a three-fold approach to research: to better detect what is living on the shelf; to better decipher and analyze the samples collected; to integrate new data with a suite of environmental variables. The goals must include a uniformity and consistency in the data presentation and more predictability in the results. Certainly a continuity of presence at the same 48 stations would enhance the knowledge and changes in community structures. Improved navigational equipment will ensure that the same site can be revisited. Yet not only should the same stations be revisited, new stations must also be analyzed In addition to box core sampling, video 81

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surveys and dive surveys would supplement the data. The hard substrates and high-relief features must be evaluated. New technologies now offer better analytical techniques. Technology might allow the tagging of certain benthic organisms to monitor movement and behavior. New technologies might provide better data for the sediment analysis including the identification and aging of constituents. Sediment traps might be used to determine sedimentation rates and the contribution by planktonic organisms. An unlimited number of questions remain unaddressed about the biology of benthic organisms and their complex interactions and environmental responses. Any additional study on life cycles, migration patterns, mobility, feeding strategies, behavior, larval recruitment and species interactions could immensely augment the level of knowledge for application to carbonate production models. Research might focus on dominant species, omitted species, recovery of live organisms, and population dynamics. The organism balances the energetic cost of calcium carbonate production against the benefits of a calcareous test. The factors affecting rate of calcification would allow better understanding of the changes in total production. Much work is also left undone on the chemical composition of these organisms and the changes in composition associated with changes in environment. Research should focus on determining the sensitivity of these assemblages to natural perturbations in the environment such as major storm events, harmful algal blooms, the green river occurrence, a major Mississippi River event, global events like El Nino or La Nina events, as well as the known anthropogenic influences of pollution, 82

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spills, net trawl damage, fishing pressure, eutrophication, dredging, changes in carbon dioxide, construction of pipelines or drilling for petroleum. Any tools that can be employed will offer better interpretation and applications of the rates estimated. Fate of Carbonate Produced The question arises, should 103 g Ca C03 m-2 yr-1 be produced across the shelf, what might the fate of this carbonate be. The carbonate on the central west Florida shelf may be broken down by a variety of agents, preserved, buried, deposited or removed from the system. With a present stand of rising sea level, is enough sediment produced and accumulated to keep up with the rise. Depositional Environment Geological events created a tectonically-stable, broad, shallow-water platform in the eastern Gulf of Mexico where marine sedimentation has occurred (Randazzo, 1997). The carbonate rock types indicate significant biological production by communities of invertebrate marine organisms (Randazzo, 1997). Historically, the sediment has been controlled by climatic settings where organic production responds to changing sea level conditions and the sediment record reflects periods of cyclic sedimentation (Randazzo, 1997). Along the coast of central west Florida, the shelf sediments consist of a 20-mile nearshore zone of quartz sand, beyond which are carbonate-dominated sediments, primarily composed of sand-sized molluscan fragments with virtually an absence of terrigenous influx into the system (MAFLA, 1975-1976; Gorsline and Swift, 1977; Sellwood, 1978; Doyle and Feldhausen, 1981; Blake and Doyle, 1983; Murray, 1991; 83

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Martinet al, 1996) The central west Florida continental shelf remains an active area of carbonate sedimentation due to this separation from siliciclastic-sediments, long-term residence in a subtropical environment, and a lack of persistent environmental stress (Hine, 1997). The thin layer of Holocene carbonate sediment is biogenic in origin with the only new source from carbonate-secreting organisms living on the shelf (Stockman et al., 1967; Gorsline and Swift, 1977; Siebold and Berger, 1982; Blake et al., 1995; Martin et al., 1996; Scott, 1997). Upon deposition, this carbonate may be subjected to many processes and conditions contributing to preservation, lithification, transport, resuspension, dissolution or formation of sediments. The benthic community both generates sediment through production and affects the rate of sediment accumulation through dissolution, transport and physical breakdown, bioerosion and bioabrasion (Cummins et al., 1986; Hallock, 1997). Carbonate may be broken down into sediment chemically by dissolution, or by mechanically breaking, sorting, abrasion or disintegration (MacGinitie and MacGinitie, 1968; Cummins et al., 1986). Many potentially preservable tests are not preserved or are poorly preserved (MacGinitie and MacGinitie, 1968; Cummins et al., 1986). No direct relationship exists for the relative age of sediments and the taphonomic alteration (Martin et al., 1996). Several decay mechanisms work to preferentially preserve organisms including: death below the sediment surface, small specimens falling into a burrow bioturbation, and reworking of the sediments by storms, currents or wave action (Cummins et al., 1986; Martin et al., 1996). Most adults are preserved while most individuals are not (Powell, 1992). 84

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The biogenic accumulation provides carbonate debris for diagenesis (MacGinitie and MacGinitie, 1968). Carbonate dissolution by microbial communities may assist in early diagenesis (Freiwald, 1995). Yet, only a small fraction of the total production will be permanently buried in the sediment (MacGinitie and MacGinitie, 1968). These coarse carbonate sediments have little mean transport or net sediment drift across or along the shelf(MAFLA, 1974; Gorsline and Swift, 1977; Doyle and Sparks; 1978). Most sediments are formed near the site of deposition (Bathurst, 1971; Murray, 1991). The Gulf of Mexico surface waters are supersaturated with calcium carbonate and well above the calcite compensation depth of 4,000 m or the lysocline at 3,000 m (Chave, 1965; MacGinitie and MacGinitie, 1968; Smith, 1971; Kennett, 1992). Thus, the carbonate-secreting organisms are neither limited by calcium carbonate nor do undersaturated surface waters dissolve them (Gladfelter, 1984). Some dissolution does occur in the interstices of the sediment at the sediment-sea water interface (MacGinitie and MacGinitie, 1968; Kennett, 1992). Death assemblages do not accumulate at the rate at which organisms die (Cummins et al., 1986). Organisms are added into the community in pulses, usually from larval recruitment events, followed by discrete pulses into the death assemblage (Cummins et al., 1986). The bottom sediments reflect this patchy nature and seasonal variations (Bishof, 1981; Doyle and Feldhausen, 1981; Cummins et al., 1986). Sedimentation Rate The central west Florida shelf is not overwhelmed by sedimentation and several estimates for the sedimentation rate of the eastern Gulf of Mexico and similar carbonate 85

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systems have been made (Murray, 1991 ) They range from 0.01 m 1000 yr1 to 2.5 m 1,000 yr1 or, 0.01 mm yr"1 to 2.5 mm yr"1 with an average rate of0.1 mm yr"1 (Moore, 1972; Blake and Doyle, 1983; Brooks, 1986; Hallock, 1986; Murray, 1991; Kennett, 1992; Powell, 1992). Approximately 101 g CaC03 m"2 yr"1 would need to be produced and accumulated to maintain a sedimentation rate of0.01 mm yr-1 At least 20% of the carbonate produced is removed through dissolution, erosion or transport from the shelf (MacGinitie and MacGinitie, 1968; Langer et al., 1997). A production rate of 102 g CaC03 yr"1 on the shelf produces a sedimentation rate of 0.1 mm yr1 And, although biological populations and sediment pulses will fluctuate, sedimentation rates linked to production rates should remain fairly constant, slow and stable (MacGinitie and MacGinitie, 1968; Gorsline and Swift, 1977; Doyle and Feldhausen, 1981; Hallock, 1981; Kennett, 1982; Blake and Doyle, 1983). The Holocene sediments are approximately 8,000 to 10,000 years old. Thus, a layer of carbonate sediments approximately one meter in depth would be expected. The thin, molluscan sand sheet shows a depth of approximately one meter in thickness deposited within the previous 8,000 to 10,000 years (Brooks, 1981; Davis, Jr., 1997; Hine, 1997). Sea Level Stand After the sea level regression in the Pleistocene, sea level has risen during the Holocene (Scott, 1997). During the late Holocene, sea level rose roughly 25 em per 100 years from 8,000 to 3,000 years before present (Davis, Jr., 1997). Since that time, the rate of sea level rise has slowed to about 4 em per 100 years or 0.4 mm yr-1 (Figure 6) (Davis, Jr., 1997). Shallow-water tropical carbonate-producing environments can keep pace with sea level rise at even faster geo-and glacioeustatic rates (Hine, 1997). With a 86

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sedimentation rate of 0 1 mm yr1 the eastern Gulf of Mexico is not keeping up with the sea level rise. Global Carbon Dioxide Budget Production of calcium carbonate is intimately connected to discussions of global carbon dioxide cycling and the classification of a carbonate-producing environment as a net source or sink of carbon dioxide. Many researchers label reefs and carbonate systems as a net source of atmospheric carbon dioxide from the one to one relationship of a mole of calcium carbonate produced to a mole of carbon dioxide released into the atmosphere (Langer et al., 1997) Others have proposed that reefs, carbonate systems and carbonate sediments are sinks for carbon dioxide through fixation by photosynthesis or calcification (MacGinitie and MacGinitie, 1968; Smith, 1978; Sundquist, 1993; Takano et al., 1994; Kayanne et al., 1995; Kleypas, 1997). Actually, both approaches are true when placed in context of time scale In terms of biological scales, diurnal changes in the carbon dioxide cycling have been noted (Le Campion-Alsumard et al., 1993; Kayanne et al., 1995). On the shorter time span of thousands of years during our current interglacial period, carbonate and reefal systems may be considered sources of atmospheric carbon dioxide (Broecker and Peng, 1987; Hallock, 1997) And, on geologic scales, limestones and carbonate environments are net sinks as they store large quantities of carbon dioxide, especially in a tectonically-stable shelf system with little transport or recycling (Hallock, 1997). To relate this to the current time frame and interest in the effects of carbon dioxide cycling on global warming, it should be noted that the estimate of carbonate produced on the central west F lorida shelf would b e n e gl igi ble to th e s tory Fossil fu e l 87

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burning has contributed more carbon dioxide into the atmosphere in the last 100 years than the reef-building communities have in 15,000 years (Hallock, 1997: Langer, et al., 1997). However, the importance of carbonate production in global cycles should not be overlooked as the benthic communities are very sensitive to environmental changes and the delicate balance between the assemblages and their environment forces their participation as mediators of climate change and responders to it (Hallock, 1997; Hubbard, 1997). Benefit of Estimated Carbonate Production Rates A picture of the calcium carbonate production rate on the central west Florida shelf concerns all who study, manage and enjoy the region and should engage the biologist, modeler, geologist, paleoceanograper, climatologist, legislator, ecologist and economist alike. The impacts of carbonate production across the eastern Gulf of Mexico are far reaching, the scope of which is nearly limitless All aspects of resource management from fisheries, to petroleum and mineral exploration and mining to the harvesting of sand are, at some point, influenced by and create impacts to carbonate production. A carbonate production rate can aid biologists in their research on living assemblages, dominant species species interactions, food supply and organisms as food supply, predation pressure, overharvesting issues, seasonality, stability, sensitivity to perturbations, effects of habitat alteration, fate of the Florida Middle Ground, potential productivity of a system, and the response to episodic events like harmful algal blooms. Of interest to the geologists are the net production sedimentation and accumulation rat e s reco r d of geoch e mical cycl es, sea l e v e l chang e res ponse and the 88

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record of evolutionary history as told by the carbonate-producing organisms. Climatologists might learn of productivity changes in relation to storm events, global climatic events and carbon dioxide cycling. A multitude of anthropogenic influences affect carbonate productivity including : pollution, runoff, spills, water quality degradation, effects of nets, trawls and dredge scars, introduction of exotic species, pipelines, the gulf as transportation, biofouling, eutrophication, creation of artificial reefs and fishery pressure. Knowledge of the consequences of these activities will greatly assist in management and directing policy for the shelf. Coastal construction and vessels increase the surface area of hard substrates for carbonate-secreting organisms. Carbonate productivity may be enhanced nearshore with these types of additions. Coastal communities continuously deal with shoreline sand issues and are constantly addressing renourishment of the beach. With all modern shelf sands of biogenic origin and thus calcium carbonate in nature, the future ofthe fine quartz beach sand is in question The sand used for renourishment likely will have a higher concentration of less desirable carbonate material and shell hash. Important to learn, as well, is at what rate the quartz sand sheet will be overcome by carbonate sedimentation. The litany of issues outlined accentuates the extent to which the carbonate productivity is integral to the health of a system and what a marvelous key to understanding these process e s it can become. The implications reach from to local, to regional to global environments 89

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Uses of Calcium Carbonate Applications utilizing calcium carbonate range from biotechnology to building to manufacturing to food production. Calcium carbonate may assist in enzyme immobilization, medicine, and ceramics and provide a source for fine chemicals (Takano et al., 1994). Calcium carbonate materials are used in building and ornamental stones, road material, cements, flux for steel smelting, lithography, optical polarizing prisms, enhancing film, pharmaceutical production like antacids and antibiotics, glass, rubber, white paint, cleaning powder and papermaking (Gaines, 1989; Burke, 1993; Callari, 1993). Calcium carbonate is found in candy, chewing gum, food fillers and toothpaste (Gaines, 1989). Therefore, demand for carbonate material is high and shortages of calcium carbonate within the world markets have been documented (Sherman, 1994). The amount produced on the central west Florida shelf is likely irrelevant, yet at some time in the future it may not be so. Production Rate Comparisons The calcium carbonate production rate estimated is compared to other shelf and carbonate producing systems in Table 16. Production rates vary from 10 to 104 g CaC03 m-2 yr-1 for the varied environments. The highest rates are found in coral-dominated habitats. Shelf production rates range from 60 to 400 g CaC03 m2 yr -1 with the central west Florida shelf production estimate of 103 g CaC03 m2 yr -1 Caution must be exercised when comparing the rate calculated in this study to those listed in Table 16 as many of the other sources were used to develop the model presented here. Therefore, trends noted might be artificial as they were created in the 90

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calculations. Having noted that, the production rate estimated appears to parallel production in other carbonate shelf systems. Comparisons do not extend beyond the central west Florida shelf. Westward of the study region approaches a different region of productivity at the shelf-slope break and beyond to the slope (Blake and Doyle, 1993). Farther north of the region the shelf is influenced by the Mississippi River and the influx of terrigenous material. South of the area, the environment changes to a tropical and more reef-dominated environment. And, west of the sites is coastal Florida with a beach system of unknown productivity 91

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Habitat CaC03 Region Dominant Source Production Producer g CaC03 m-2 yr-1 Temperate Shelf 400 California Borderland Foraminifera Smith (1970, 1971) Gross Production 104 Hypothetical Reef Models Coral Chave et al. (1972) Net P r oduc t ion 10 3 Sublittoral 1 400 Biscayne, Florida Macrobenthos Moore (1972) Intertidal 1 X 10 3 Reefs 4 X 10 3 Coral Smith and Kinsey Protected Lagoons 8 X 102 (1976) ShelfEdge Reef System 1.2 X 10 3 St. Croix, U.S. Virgin Coral Hubbard et al. (1988) Islands Bank Area 331 Buchanan Bank Porites, Thalassia Bosence (1989) Reef Area 1 X 10 3 Upper Cross Bank Thalassia epibionts Non-Carbonate Shelves 25 Milliman (1993) Banks and Bays 500 Coral Reefs 1.5 X 10 3 Halimeda Bioherms 3 X 10 3 Carbonate Shelf 60 Milliman (1993) Phytobenthic 100 Mallorca-Menorca Shelf Algal dominated Canals and Ballesteros Communities NW Mediterranean (1997) Low Productivity 1.2-120 Coral Reefs Foraminifera Langer et al. ( 1997) High Productivity 30-1000 Carbonate Shelf 103 Central West Florida Shelf Mollusca This Study 25-434

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Concluding Remarks The goal of this study was to describe the benthic carbonate-producing communities and to quantify the calcium carbonate production rate for the central west Florida shelf. Benthic communities represent good indicator organisms, both as bioindicator and geoindicator (Blake et al., 1995; Lidz and Hallock, 2000). They provide a basis upon which to monitor ecosystems, note the reality of environmental and anthropogenic perturbations and assess living and relict communities (Blake et al., 1995; Lidz and Hallock, 2000) The previous discussion has demonstrated the complexity of this type of system and, with little data available and incomplete research, the difficulty in developing a production rate. Non-linear biological systems demand a complete look at many variables, both in space and in time. This preliminary work provides the foundation for a dynamic working model for quantitatively representing production rates on the shelf. Using extensive though not comprehensive data from assemblages found on the central west Florida shelf, a model for calcium carbonate production was designed and presented. Few patterns have been noted for production rates as a function of depth, latitude or biotopes. However, substrate may play the major role in community distribution, composition, larval recruitment and productivity on the shelf (MAFLA, 1974; Babashoff, 1982; Blake and Doyle, 1983; Blake et al., 1995; Canals and Ballesteros, 1997). The benthic community thus both shapes the substrate and is shaped by it. Understanding these interactions will aid in refining the production processes and rates for the shelf 93

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The assemblages and community composition varied temporally yet the production values did not vary dramatically. The short-term changes of pulsating patches do not appear to affect long-term stability of the system nor the stability of the production rates. Although the communities vary spatially across the shelf and reside in dissimilar hydro-biological zones, the production rates for the stations are similar. To illustrate, compare the production rates for the Florida Middle Ground at 114 g CaC03 m-2 yr-1 and the surrounding shelf estimated to be 101 g CaC03 m-2 yr-1 Production rates are quite close in value but the biotope and populations of organisms within the assemblage varied greatly and secreted carbonate at different rates. This ecologically rich shelf environment shows remarkable stability while still sensitive to perturbations (MAFLA, 1975-1976). The results reflect healthy, responsive benthic communities producing calcium carbonate at a rate of 102 g CaC03 m-2 yr-1 94

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APPENDICES 106

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Appendix 1: List of Stations on the Central West Florida Shelf Station MAFLNMMS Transect Depth Longitude Latitude Date of No. Sta. No. inm Sample 1 42 II 37 -842630 28 41 59 6/9/1974 2 43 II 45 -842800 28 30 00 6/14/1974 3 44 II 44 -842321 28 26 29 6/15/1974 4 45 II 53 -842359 28 21 00 6/15/1974 5 46 II 37 -842001 28 41 59 6/9/1974 6 47 II 36 -842012 28 34 00 6/10/1974 7 48 II 40 -842100 28 29 00 6/14/1974 8 49 II 42 -842100 28 24 00 6/15/1974 9 50 II 48 -842058 28 19 00 6/15/1974 10 52 II 54 -841732 28 13 59 6/15/1974 11 53 II 37 -841301 28 41 59 6/10/1974 12 54 II 34 -841059 28 29 00 6/10/1974 13 55 I 44 -835256 27 56 33 6/14/1974 14 56 I 38 -834449 28 00 38 6/16/1974 15 57 I 37 -834229 27 57 30 6/16/1974 16 58 I 43 -834132 27 47 58 6/13/1974 17 60 I 31 -833530 28 01 00 6/16/1974 18 61 I 33 -833356 27 52 31 6/17/1974 19 62 I 34 -833059 27 50 01 6117/1974 20 63 I 30 -832729 27 56 00 6/17/1974 21 64 I 30 -832500 27 50 00 6/1711974 22 65 I 42 -832530 27 45 30 6/13/1974 23 2101 I 11 -821508 26 24 59.6 6/75-2/76 24 2102 I 18 -822459.6 26 24 59.6 6/75-2/76 25 2103 I 37 -825759.7 26 25 00.0 6/75-2/76 26 2104 I 53 -832300.8 26 25 00.0 6/75-2/76 27 2105 I 90 -834957.6 26 24 59.5 6/75-2/76 28 2106 I 168 -841500 26 24 56.8 6175-2176 29 2207 II 19 -830900.3 27 57 00.4 6/75-2/76 30 2208 II 31 -832729.6 27 56 00.5 6/75-2/76 31 2209 II 34 -833359 27 52 30.5 6/75-2/76 32 2210 II 37 -834229.2 27 57 28.8 6/75-2/76 33 2211 II 43 -835259.5 27 56 29.5 6/75-2/76 34 2212 II 189 -844759.6 27 57 00.0 6/75-2/76 107

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Appendix 1: (Continued) Station MAFLA/MMS Transect Depth Longitude Latitude Date of No. Sta. No. inm Sample 35 2313 m 176 -851503 28 23 59.3 6/75-2/76 36 2314 m 29 -842059 28 29 58.6 6/75-2/76 37 2315 m 38 -842009.1 28 33 59.1 6/75-2/76 38 2316 m 35 -842000.7 28 42 00.3 6/75-2/76 39 2317 m 29 -840559.9 28 56 00.3 6/75-2/76 40 2318 m 20 -834500.5 29 05 00.8 6/75-2/76 41 IC I 13 -824945 273791 7/92-5/94 42 ID I 13 -824945 273791 7/92-5/94 43 nc n 6 -823562 271562 7/92-5/94 44 llD n 6 -823562 271562 7/92-5/94 45 me m 6 -822588 265966 7/14/1992 46 IDD m 6 -822588 265966 7/14/1992 47 IVC IV 6 -824180 272627 10/29/1993 48 IVD IV 6 -824180 272627 10/29/1993 108

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Appendix 2: Large Benthic Foraminiferal Production 1974 Transect II STATION NO. I 2 3 4 5 6 7 8 9 10 II 12 AVGS. DEPTII (METERS) 37 45 44 53 37 36 40 42 48 54 37 34 42 PLANKTIC!BENTIIIC RA 110 6:100 9:100 8:100 14:100 3:100 4:100 5:100 6 : 100 5:100 10:100 2:100 5:100 % LIVING BENTIIICS 9% 15% 14% 36% 13% 19% 13% 13% 12% 14% 15% 10% 15% LNING SPECIMENS/SAMPLE 1527 446 278 817 414 191 1158 453 481 2920 508 158 779 LIVING SPECIMENS/m2 x 104 310.93 90.77 56.69 166.40 84.32 38.90 235 .91 92.19 97 90 594 .7 0 103.53 32.11 158. 70 %CARBONATE PRODUCERS 93% 86% 77% 88% 87% 75% 83% 77% 79% 87% 86% 80% 83% CARBONATE PRODUCERS 289.48 77.97 43.88 146.59 73 69 28 98 194. 86 70 62 77.34 519 77 88.73 25 69 136.47 in No. I m 2 x 10 4 Amphistegina gibbosa 8.17 3.40 0.50 2.28 1.83 6.37 3.69 13. 90 2.80 1.38 3.69 CARBONATE PRODUCTION RATE in g CaCO 3/m 2 lyr 0.00 27 77 11. 56 1.70 7 74 6 22 21. 66 12.54 47.26 0.00 9.50 4 69 12.55 0 \0 Archaias angulatus 1.66 0.25 0.16 Sorites hofkeri 3.33 3 63 0.58 SUBTOTAL 4.99 3.88 0 .74 CARBONATE PRODUCTION RATE in g CaCO 3/m 2 lyr 5.99 4 .65 0.89 Peneroplis carinatus 3.11 6.63 0 .57 6.16 2 53 1.28 0 .65 0.29 7.56 2.67 2.62 Penerop/is proteus 0.27 SUBTOTAL 3.11 6 90 0 57 6.16 2.53 1.28 0.00 0.65 0.29 0 00 7.56 2.67 2.64 CARBONATE PRODUCTION RATE in g CaCO 3/m 2 lyr 3.73 8.28 0 68 7.39 3.04 1.54 0.00 0 .77 0.35 0 00 9.07 3 20 3.17 TOTAL CaC03 PRODUCTION RATE 3.73 36 05 12.24 15. 08 15. 43 7.76 21.66 13.31 47.62 0.00 18.57 7.89 16.61

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Appendix 3 : Large Benthic Foraminiferal Production 1974 Transect I STATION NO. 13 14 15 16 17 18 19 20 21 22 AVGS DEPTH (METERS) 44 38 37 43 31 33 34 30 30 42 36 PLANKTIC/BENTHIC RATIO 12:16 0 3:100 4:100 4:100 2:100 1 : 100 1 : 100 2:100 0:100 1:100 % LIVING BENTHICS 16% 12% 6 % 12% 15% 6% 13% 8% 7 % 10% 11% NO. LIVING SPECIMENS/m2 x 104 467.21 295 66 242.57 42.23 642.43 372.84 63.00 309.57 235 98 469.52 314.10 %CARBONATE PRODUCERS 87% 94% 93 % 86% 85% 85% 95% 86% 93% 85% 89% CARBONATE PRODUCERS 408.34 278.51 225.59 36.15 547 99 316.55 59.91 265.61 218.99 400.03 275.77 in No. /m 2 X /04 Amphistegina gibbosa 4 67 7. 08 1.18 Gypsina vesicularis 1.93 0.19 SUBTOTAL 4 67 1.93 7 08 1.37 CARBONATE PRODUCTION RATE in g CaCO 3/m 2 lyr 15 89 6.55 60.18 8.26 -0 Archaias angulatus 4.7 0 0.4 7 Sorites hojkeri 3.27 8.87 1.70 1 7.35 5.26 8.73 1 0.80 5 60 SUBTOTAL 3.27 8.87 1.70 17.35 5 .26 8 7 3 15.49 6 07 CARBONATE PRODUCTION RATE in g CaCO 3/ m 2 lyr 3 92 10.64 2.04 20.81 15. 79 26.19 18 59 9.80 P e n e ropli s carinatus 1.40 5.91 8 .35 6 19 7.98 2 98 P e neroplis prot e us 1.41 0 14 SUBTOTAL 1.40 5 .91 8.35 6.19 9 .39 3.12 CARBONATE PRODUCTION RATE in g CaCO 3/m 2 /yr 1.68 7.10 10 02 7 .4 3 11. 27 3.7 5 TOTAL LARGE FORAMINIFERA 9.34 14.78 1.70 0.00 27.62 0.00 0 00 11.45 15.8 1 24.88 10.56 %OF CARBONATE PRODUCERS 2% 5% 1% 0% 5% 0% 0% 4% 7% 6% 3% TOTAL CaC03 PRODUCTION RATE 21.49 1 7.7 4 2.04 0 00 37 .39 0.00 0 00 23 22 86.3 7 29 86 21.8 1

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...... ...... ...... Appendix 4: Large Benthic Foraminiferal Production Summer 1975 Transects I and II STATION NO. 23 24 25 26 27 28 I 29 30 31 32 AVG. DEPTH (METERS) 11 18 37 53 90 168 63 19 31 34 37 % LIVING BENTHICS 2 90 11.50 6 00 12.30 4.40 17. 20 9.05 12 80 12.60 7 50 14.10 T O TAL LIVING/m2 x 105 1.67 0.748 0.95 3.48 1.71 28.35 6.15 13.2 5 38.10 22.70 3.74 %CARBONATE PRODUCERS 88% 96% 95% 82% 83% 94% 90% 98% 89% 89% 86% CARB O NATE PRODUCERS in No. l m 2 X /05 1.46 0.718 0 90 2.85 1.42 26 .64 5.67 13.03 34.04 20 .12 3.22 Amphistegina gibbosa 0 12 0.09 1.32 0.25 0 .01 CARBONATE PRODUCTION RATE in g CaCO Jim 2 lyr 4 03 2.90 44.98 8.65 0.42 Archaias angulatus O.Ql 0.017 0.00 0.13 Sorites hofkeri orbitolitoides 0.008 O.Ql 0.00 0 .38 0.45 O.Ql SUBTOTAL O.Ql 0.025 0.01 O.Ql 0.51 0 .4 5 O.Ql CARBONATE PRODUCTION RATE in g CaCO Jim 2 lyr 0.17 0.752 0 07 0 16 6.10 5.45 0.15 Peneroplis bradyi 0.01 0.00 Peneroplis carinatus 0.004 0.04 0 .13 0.09 0.04 0.15 0.05 Peneroplis proteus 0.00 SUBTOTAL 0.004 0.04 0 .14 0.09 0.05 0 .15 0.05 CARBONATE PRODUCTION RATE in g CaCO Jim 2 lyr 0 05 0.45 1.70 1.13 0.56 1.82 0.60 CARBONATE PRODUCTION RATE BY LARGE BENTHICS Summer 1975 in g CaCO 3Im 2 lyr 0 .17 0.802 0.45 5.73 2.97 46.11 9.37 0.00 6.10 7.26 1.17 33 34 II AVG. 43 189 59 13.90 27 80 14.7 8 1.99 45.06 20 .81 85% 98% 91% 1.70 44 .31 19.40 0.07 0.01 2.25 0.45 0.02 0.15 0.17 0.15 0 1 9 1.80 2.25 0.00 0.03 0.00 0.03 0.40 2.25 1.80 3 .10

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Appendix 5: Large Benthic Foraminiferal Production Summer 1975 Transect III and Averages STATION NO. 35 36 37 38 39 40 III SUMMER 1975 AVG AVERAGE DEPTH (METERS) 176 43 38 35 29 20 57 60 % LIVING BENTHICS 37.60 2 .60 10.60 21.90 12.12 11.98 TOTAL LIVING/m2 x 10 5 29.89 23.59 3.89 10.39 33 .31 5 .67 17. 79 14 92 %CARBONATE PRODUCERS 99% 90% 90% 90% 92% 98% 93% 91% CARBONATE PRODUCERS in No 1m2 x 105 29.69 21.15 3.49 9.32 30. 54 5.58 16.63 13. 90 Amphistegina gibbosa 0 10 0.31 0.05 0 14 0.10 0 .12 CARBONATE PRODUCTION RA T E in g CaCO 3/m 2 lyr 3 39 10.69 1.76 4.71 3.43 4.1 7 --N Archaias angulatus 0.04 0.01 0 .01 Sorites hojkeri orbitolitoides 0 22 0.06 0.05 0 07 SUBTOTAL 0 22 0 09 0.05 0 08 CARBONATE PRODUCTION RATE in g CaCO 3/m 2 lyr 6 64 2.84 1.58 1.33 Peneroplis bradyi 0 .11 0 02 0.01 Peneroplis carinatus 0 16 0.03 0 07 0.44 0.04 0.12 0.07 Peneroplis proteus 0.08 0.01 0 .03 0.11 0.02 0 .04 0.01 SUBTOTAL 0 .24 0 .04 0.10 0.66 0.06 0.18 0 09 CARBONATE PRODUCTION RATE in g CaCO 3/m 2 lyr 2.83 0.47 1.25 7.97 0 68 2 20 1.05 CARBONATE PRODUCTION RATE BY LARGE BENTHICS Summer 1975 in g CaCO 3/m 2 lyr 3.39 13. 52 2.23 5 96 14.61 3.52 7 20 6.56

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Appendix 6: Large Benthic Foraminiferal Production Fal11975 Transects I and II STATION NO. 23 24 25 26 27 28 I 29 30 31 32 33 34 II AVG. AVG. DEPTH (METERS) 11 18 37 53 90 168 63 19 31 34 37 43 189 59 % LIVING BENTHICS 15 10 13.40 10.80 6.50 11.60 22.70 13.35 12 10 13.30 19.70 17 10 1 1.70 20 .60 1 5.75 TOTAL LIVING/m2 x 10 5 30.94 17.66 13.04 2.17 11.84 26.04 16.95 15.10 43 73 70.57 30.18 11.72 10.24 30.26 %CARBONATE PRODUCERS 87.67 96.09 95 .15 81.97 83.33 94.00 89.70 98.33 89.33 88.67 86.29 85.33 98.33 91.05 CARBONATE PRODUCERS in No I m 2 x 10 5 27 .13 16.97 12.40 1.78 9.87 24 47 15.44 14.85 39.06 62.57 26.04 10.00 10.07 27.10 Amphistegina gibbosa 0.07 0.59 1.22 0.31 0.10 0.39 0.08 CARBONATE PRODUCTION in g CaCO Jim 1 lyr 2.51 20.13 41 .31 1 0.66 3.43 13.29 2.79 Archaias angulatus 0.10 0.39 0.08 0.15 0.02 Sorites hofkeri orbitolitoides 0.20 0.04 0.04 0.44 1.41 0.10 0.03 0.33 SUBTOTAL 0.10 0.59 0.04 0.12 0.58 1.41 0.10 0.03 0.35 CARBONATE PRODUCTION in g CaCO Jim 2 lyr 3.09 17 .76 0.47 3 55 7.00 16.94 1.21 0.41 4.26 Peneroplis bradyi 0 .01 0.00 0.00 Peneroplis carinatus 0.10 0 52 0.08 0 09 0.13 0.47 0.40 0.15 Peneroplis proteus 0.00 SUBTOTAL 0.10 0.52 0.09 0.09 0.13 0.47 0.40 0.15 CARBONATE PRODUCTION in g CaCO Jim 1 lyr 1.18 6.20 1.06 1.04 1.58 5.65 4.85 1.75 CARBONATE PRODUCTION RATE BY LARGE BENTHICS Fall 1975 in g CaCO Jim 2 lyr 3.09 18.94 6 .20 3.57 20.61 42.35 15.80 0.00 7.00 22.58 9.49 13.29 0.41 8.79

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Appendix 7: Large Benthic Foraminiferal Production Fall1975 Transect III and Averages STATION NO. 35 36 37 38 39 40 III FALL 1975 AVG. AVG. AVERAGE DEPTH (METERS) 176 43 38 35 29 20 57 60 60 % LIVING BENTHICS 39.30 14.10 21.70 10.50 11.30 49.60 24.42 17.84 17.84 TOTAL LIVING/m2 x 10 5 63.63 5.43 54.36 1.61 4 66 6.40 22.68 23. 29 23.29 %CARBONATE PRODUCERS 99.33 89.67 89.67 89 67 91.69 98.33 93.06 91.27 91.27 CARBONATE PRODUCERS in No. 1m2 x 105 63.20 4.87 48.74 1.44 4.27 6.29 21.47 21.34 21.34 Amphistegina gibbosa 0.21 0.02 0.18 0.02 0.07 0 .16 0.16 CARBONATE PRODUCTION in g CaCO 3/m 2 lyr 7.21 0.62 6.16 0 .73 2.45 5 30 5.30 Archaias angulatus 0.04 0 .01 0 04 0 04 Sorites hojkeri orbitolitoides 0.03 0 06 0.02 0 .13 0.13 SUBTOTAL 0.03 0.11 0.02 0.17 0.17 CARBONATE PRODUCTION in g CaCO 3/m 2/y r 0.93 3.20 0.69 2 .83 2.83 Peneroplis bradyi 0.02 0.00 0 00 0.00 P enerop lis carinatus 0.04 0.36 O.Ql 0.06 0.04 0 09 0.12 0.12 Peneroplis proteus 0.02 0.18 O.Ql 0.02 0.02 0.04 O.ot O.Ql SUBTOTAL 0.05 0.54 0 02 0.09 0.06 0.13 0.14 0.14 CARBONATE PRODUCTION in g CaCO 3/m 2 /yr 0.65 6 52 0.19 1.11 0.77 1.54 1.62 1.62 CARBONATE PRODUCTION RATE BY LARGE BENTHJCS Fal/1975 in g CaCO 3/m 2 !yr 7.21 1.27 12.68 0.92 2.04 3.97 4.68 9.76 9.76

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Appendix 8: Large Benthic Foraminiferal Production Winter 1976 Transects I and II STATION NO. 23 24 25 26 27 28 I 29 30 31 32 33 34 II AVG. AVG. DEPTH (METERS) 11 18 37 53 90 168 63 19 31 34 37 43 189 59 % LIVING BENTHICS i5.4 17.8 6.7 18.8 6.9 9.7 12.6 9.5 9.7 9.1 16.0 13.4 24.3 13.7 TOTALLIVING/m2 x 105 4.28 11.13 11.42 15.81 2.24 6.94 8.63 6.96 15.63 14.77 9.04 2.36 6.27 9.17 %CARBONATE PRODUCERS 87.67 96.09 95.15 81.97 83.33 94.00 89.70 98.33 89.33 88.67 86.29 85.33 98.33 91.05 CARBONATE PRODUCERS in No. / m 2 X J05 3.75 10.69 10.86 12.96 1.86 6.52 7 .77 6.85 13.96 13.10 7.80 2.01 6.16 8.31 Amphistegina gibbosa 0.54 0.11 0.32 0.16 0.03 0.08 0.02 CARBONATE PRODUCTION RATE -ing CaCO 31m2 /yr 18.28 3.80 11.00 5.51 1.03 2.67 0.62 VI Archaias angulatus 0.01 0.25 0.04 0.05 Sorites hofkeri orbitolitoides 0.12 0.01 0.02 0.16 0.30 0.03 0.02 0.08 SUBTOTAL 0.01 0.37 0.01 0.07 0.21 0.30 0.03 0.02 0.09 CARBONATE PRODUCTION RATE in g CaCO 3/m 2 /yr 0.43 11.19 0.09 1.95 2.50 3.54 0.36 0.25 1.11 0.00 Peneroplis bradyi 0.05 0.01 0.00 Peneroplis carinatus 0.06 0.45 0.59 0.02 0.19 0.10 0.12 0.04 Penerop/is proteus 0.00 0.00 SUBTOTAL 0.06 0.45 0.65 0.02 0.20 0.10 0.12 0.04 CARBONATE PRODUCTION RATE in g CaCO 3/m 2 lyr 0.75 5.43 7.74 0.28 2.37 1.18 1.45 0.44 CARBONATE PRODUCTION RATE BY LARGE BENTHICS Winter 1975 in g CaCO 3/m 2 lyr 0.43 11.93 5.43 26.03 3.89 11.28 9.83 0.00 2.50 4.73 2.84 2.67 0.25 2.17

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Appendix 9: Large Benthic Foraminiferal Production Winter 1976 Transects III and Averages STATION NO. 35 36 37 38 39 40 III-FMG WINTER 1 976 AVG. AVERAGE D EPTH (METERS) 176 43 38 35 29 20 57 60 % LIVING BENTHICS 27.3 30.8 19.6 16. 3 8.2 40 1 23 7 1 6 6 TOTAL L IVING/m2 x 10 5 41. 6 1 5 70 4 .21 0 72 16.50 1.44 1 1.70 9. 8 3 %CARBONATE PRODUCERS 99 33 89 67 89 67 89 67 91.69 98 .3 3 93 .06 91.27 CARBONATE PRODUCERS in No./m 2 X /05 41.33 5.11 3.78 0.64 15.13 1.42 11.2 3 9.11 Amphi s tegina gibbosa 0.14 0 02 0 .01 0.01 0.03 0 .07 CARBONATE PRODUCTION RATE in g CaCO 3/m 2 /yr 4.72 0.65 0.48 0.33 1.03 2. 3 9 -0\ Archaia s angulatus 0.01 0.00 0 02 Sorit e s hojkeri orbitolitoides 0.11 0.01 0.02 0 .04 S U BTOTAL 0.11 0 02 0.02 0 06 CARBONATE PRODUCTION RATE in g CaCO 3/m 2/yr 3.29 0.72 0.67 1.2 4 Peneroplis bradyi 0.05 0.01 0.01 Peneroplis carinatus 0 00 0.22 0.01 0.04 0 09 Peneroplis proteus 0 .00 0 05 0 00 0.01 0 .0 0 SUBTOTAL 0.0 1 0.33 0.0 1 0.06 0.1 0 CARBONATE PRODUCTION RATE in g CaCO 3/m 2 lyr 0 09 3.95 0.17 0.70 1.17 CARBONATE PRODUCTION RATE BY LARGE BENTHICS W int er 1975 in g CaCO 3/m 2 /yr 4.72 0 65 0.48 0 .4 1 7 24 0 90 2.40 4. 80

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Appendix 10: Small Benthic Foraminiferal Production 1974 Transect II STATION NO. 1 2 3 4 5 6 7 8 9 10 11 12 AVG DEPTH (METERS) 37 45 44 53 37 36 40 42 48 54 37 34 42 PLANKTIC/BENTHIC RATIO 6:100 9:100 8:100 14:100 3 : 100 4:100 5:100 6:100 5:100 10:100 2:100 5:100 % LIVING BENTHICS 9.2 15 13.6 36.1 12.5 18.7 12.9 13.2 12.2 13.9 15.2 10.1 15 LIVING SPECIMENS/CM2 310.93 90.77 56.69 166.40 84.32 38.90 235.91 92.19 97.90 594.70 103.53 32.11 159 %CARBONATE PRODUCERS 93.1 85.9 77.4 88.1 87.4 74.5 82.6 76.6 79 87.4 85.7 80 83 CARBONATE PRODUCERS 289.48 77.97 43.88 146.59 73.69 28.98 194.86 70.62 77.34 519.77 88.73 25.69 1 36 Ammonia beccarii parkinsoniana 0.50 0.29 0 Ammonia translucens 0.27 0 Articulina mucronata 0.12 0 Articulina pacifica 0.27 0.10 0 Articu/ina 0.50 0 Asterigerina carinata 1.82 0.74 10.48 5.87 7.97 5 Brizalina fragi/is 0.50 0 Brizalina /owmani 2.44 2.16 0.89 1.27 13.68 2.38 4 Cancris oblonga 2.09 2.44 0.50 2 Cancris sagra 1.54 1.16 2.36 0.69 1 Carterina spiculotesta 0.17 0 Cassidulina crassa 0.98 1 Cassidulina curvata 2.16 0.65 0.29 1 Cassidulina subglobosa 33.27 7.90 1.87 6.66 2.78 1.67 19.58 3.41 1.96 25.57 6.94 10 Cibicidesfloridanus 13.37 9.71 8.11 22.13 4.47 0.66 13.45 11.34 13.02 47.58 7.97 2.15 13 -Cibicides io 0.91 0.69 1 Cibicides mol/is 1.16 1 Cibicides :.p. 0.17 0 Discorbis sp. 2.16 2 Elphidium advenum 0 50 0 Elphidium delicatulum 0.96 1 Elphidium discoidale 6.22 0.40 1.27 19.63 7 Eponides antillarum 3.33 1.43 0.98 13.68 5 Fissurina :.pp. 0.50 0 Fursenkoina mexicana 1.82 1.94 0.69 10.11 4

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Appendix 10: (Continued) Fursenkoina pontoni 0.9I 1.13 2.83 2.28 7.08 2.49 0.69 0.87 2 Glabratella ? sp. 0.29 0 Guttu/ina australis 0.84 I Guttu/ina laevis 1.16 0.98 I Hanzawaia strattoni 32.03 3.90 1.30 28.29 I2.39 1.17 26.66 4.89 6.56 136.78 20.40 3.2I 23 Lenticulina orbicularis 0.69 I Loxostomum truncatum 0.7I I Marginu/ina planata 0.29 0 Miliolinel/a circularis 5.29 0.64 1.13 2.83 2.28 2.06 6.37 3.69 0.69 5.95 0.22 3 Miliolinella sp. Neoconorbina orbicularis 9.08 5.67 6.I6 7.34 4.67 37.04 4.89 4.2I II7.I6 I4.49 5.88 20 Nodobaculariella cassis 1.66 0.25 0.29 I Nonion depressulum 0.27 0.74 0.50 0.84 0.7I I matagordanum Nonionel/a atlantica 0.93 O.I7 0.50 4.22 1.56 0.7I 2.I2 0.29 13.68 3 -00 Oo/ina melo 0.17 0 Planorbulina mediterranensis I4.6I 1.66 2.28 1.95 8.73 II.34 0.69 6 Planulina exorna 8.40 3.36 3.57 8.82 6.49 1.67 IO.I4 5.53 4.60 7.73 0.96 6 Poroeponides latera/is 0.50 1.10 1.17 1.66 I Pseudonodosaria mayori O.I2 0 Pyrgo nasutus 3.83 0.39 2 Pyrgo subsphaerica 0.93 O.I2 I Quing uelocu/ina bicostata 5.29 5 Quinquelocu/ina bicarinata 0.50 0 Quinquelocu/ina bosciana 0.50 0.59 0.27 0 Quinqueloculina cultrata 1.16 I Quinquelocu/ina horrida 0.29 0 Quinquelocu/ina lamarckiana I20.33 5.I7 4.36 3.83 6.75 1.44 I7.22 2.49 0.69 61.25 I0.04 2.99 20 Quinquelocu/ina la evigata 1.65 2 Quinqueloculina polygona 0.50 0 Quinqueloculina seminulum 1.78 0.3I I IQ_uinquelocu/ina subpoeyana 1.16 I

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Appendix 10: (Continued) Quinqueloculina spp. 0.91 2.16 0.25 0.39 0.71 0.29 1.78 1 Reussella atlantica 5.29 2.72 2.10 2.83 6.37 1.84 2.64 0.55 3 Rosa/ina columbiensis 31.09 7.26 1.53 3.33 4.22 4.55 25.95 10.42 6.85 39.85 7.56 3.76 12 Rosa/ina concinna 0.29 0 Rosa/ina jloridana 0.27 3.33 0.39 1 Rosa/ina jloridensis 0.27 0.69 0 Rosa/ina suezensis 0.50 0 Rosa/ina sp 0.17 0.29 0 Sawina pulchella primitiva 0.17 0.29 0 Sigmoilina distorta Siphonina pu/chra 0.29 0 Spirillina vivipara 0.12 0 Spiroloculina wata 0.28 1.78 1 Spiroloculina soldanii 0.27 0.50 0 Spiroloculina sp. 0.71 1 Tretomphalus planus Trlfarina bella 0.50 0.29 0 Trifarina bradyi 0.29 0 Trilarina Jamaicensis 0.12 0 Triloculina brevidentata 0.71 1 Triloculina rotunda 0.50 0 Triloculina tricarinata Triloculina trigonula 0.40 0.50 0.31 0 Triloculina sp. 1.78 2 Wiesnerella auriculata 9.33 1.54 2.28 0.39 1.65 0.92 0.96 2 TOTAL SPECIMENS 286.37 62.90 39.91 134.95 65.01 25.87 188.49 66.29 63.15 519 . 77 78.38 21.64 129 %OF CARBONATE PRODUCERS 98.93 80.68 90.96 92.05 88.22 89.26 96.73 93.86 81.66 100.00 88.34 84.25 90 NO. SPECIES 14 23 23 42 20 22 20 15 37 17 10 11 21 % OF NO. 0 SPECIES 93.33 88.46 92.00 91.30 83.33 91.67 95.24 88.24 94.87 100.00 83.33 84.62 91 TOTAL CARBONATE PRODUCTION ingCaC03 m -2 yr I 47.46 10.42 6.61 22.36 10.77 4.29 31.24 10.99 10.465 86.14 12.99 3.59 21

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...... N 0 Appendix 11: Small Benthic Foraminiferal Production 1974 Transect I STATION NO. 13 14 15 16 17 18 19 20 DEPTH (METERS) 44 38 37 43 31 33 34 30 PLANKTIC/BENTHIC RATIO 12:160 3:100 4:100 4:100 2:100 1:100 1:100 2:100 % LIVING BENTHICS 16.3 11.5 6.2 12 15. 1 6.2 13.3 8.3 NO. LIVING SPECIMENS/CM2 SEDIMENT 467.21 295.66 242.57 42.23 642.43 372.84 63.00 309.57 %OF CARBONATE PRODUCERS 87.40 94.20 93.00 85. 60 85.30 84.90 95.10 85.80 CARBONATE PRODUCERS 408.34 278.51 225.59 36.15 547.99 316.55 59.91 265 .61 Ammonia beccarii tepida 5.91 0.30 1.93 0.19 Amphicoryne scalaris 0.89 Asterigerina carina/a 1.40 1.70 0.42 14.78 11.78 Brizalina fragilis 1.40 Brizalina goessii 1.40 Bri zalina lowmani 1.40 1.93 Bri za lina sp Bulimin e lla e l ega ntissima 1.40 Cancris oblonga 7.94 Cancris sagra 3.27 1.93 Cassidulina subglobosa 7.94 19.81 3.15 Cibicides floridanus 34.11 7.98 21.10 5.19 59.75 24.98 Cibicides io 4.50 Cibicides mollis 9.34 Cibicides sp. Cribroe/phidium poeyanum 6.42 Discorbis sp. 10.75 Elphidium advenum 0.89 Elphidium delicatu/um 10.75 0.19 Elphidium discoidale 4.67 9.76 5.58 3.08 19.27 12.30 Elphidium galvestonense 3.27 Eponides antillarum 10.75 2.43 10.92 7.12 Fissurina s pp. 4.67 Fursenkoina comp/anata 1.40 0.89 Fursenkoina mexicana 3.27 2.07 Fursenkoina pon toni 40.65 3.84 10.92 21 22 AVG 30 42 36 0:100 1:100 7 1 9.7 235.98 469.52 314.10 92.80 85.20 88.93 218 99 400.03 275.77 3.29 1.16 0 09 32.87 6.29 0.14 0 .14 1.41 0.47 1.41 0.14 0.14 0.79 1.41 0.66 3.09 51.65 20.48 0.71 0.52 3.29 1.26 1.41 0.14 6.10 1.25 1.07 1.41 0.23 1.09 31.46 8.61 1.41 0.47 6.37 9.39 4.70 0.47 1.41 0 37 1.41 0.67 7.98 6.34

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Appendix 11: (Continued) Fursenkoina pontoni 40 65 3 84 10 92 7.98 6 34 Fursenkoina spinicostata 4 67 0.47 Guttulina australis 3 2 7 6 .4 2 4 .70 1.44 Guttulina hirsuta 1.93 0.19 Guttulina laevis 3.27 6.42 0.93 4.70 1.5 3 Hanzawaia strattoni 49 99 79.83 58.94 4.22 100.86 120.43 8 .19 103.09 54. 28 87 80 66. 7 6 Lagena spp 1.40 0.14 Miliolinella circularis 1.40 1 3.90 11.40 1.14 1.93 0 19 1.41 3 .14 Neoconorbina orbicularis 35. 98 16.98 6 33 66.17 12. 30 8 82 13.31 16.52 26.76 20.32 Nodobaculariella cassis 1.40 4.50 1.41 0.73 Nonion depressulum matagordanum 3.27 0.93 0.42 Nonionella atlantica 12 .61 5. 0 3 0 72 19. 27 11.19 2 .71 8.36 6.10 6.60 Pavonina atlant i ca 1.40 0 1 4 Planorbulina mediterranensis 15.42 16.25 8.3 5 10 .15 5 02 Planulina exorna 31.30 8.00 14.78 11.19 1.4 1 6.67 ...... N Poroeponides latera/is 3.27 0.33 ...... Pseudonodosaria mayori 0.89 0 09 Pyrgo dent i culata 1.41 0 .14 Pyrgo depressa 0.71 0 0 7 Pyrgo nasutus 6.07 0.42 1.93 1.12 3 10 4. 70 1.7 3 Quinqu e loculina bicostata 4.50 0.71 1.41 0.66 Quinqueloculina bicarinata 1.93 0.19 Quinqueloculina bosciana 0 .71 0.0 7 Quinqueloculina cultrata 1.40 0.14 Quinqueloculina dutemplei 1.41 0 .14 Quinqueloculina lamarckiana 7.94 91.65 53 36 6.46 4.50 84.64 8 82 40.24 59 00 20.19 37.68 Quinqu eloculina /aevigata 0.89 0 .13 1.93 0 .19 0 .71 0 38 Quinqu e loculina polygona 0.13 1.12 2.17 0.3 4 Quinqueloculina s e minulum 1.93 1.41 0.33 Quinqueloculina subpoeyana 3.2 7 1.12 0 .44 Quinqueloculina spp. 3 2 7 0 93 0.42 Reussella atlantica 17.29 0 30 25.70 1.70 4 70 4 97

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...... N N Rosa/ina bulbosa Rosa/ina columbiensis Rosa/ina concinna Rosa/ina jloridana Rosa/ina jloridensis Rosa/ina suezensis Rosa/ina sp. Sagrina pu/chella primitiva Spiroloculina sp. Trifarina bella Trilocu/ina brevidenta Triloculina trigonula Triculina sp. Valvulineria mexicana Wiesnerella auriculata TOTALSMALLCARBONATEPRODUCERS %OF TOTAL CARBONATE PRODUCERS NO. SPECIES TOTAL CARBONATE PRODUCTION ingCaC03 m .J yr Appendix 11: (Continued) 0.13 18.63 24.98 6.33 59.75 32.12 14.02 8.35 0.89 3.27 4.50 4.67 0.30 6.42 1.93 1.40 0.55 1.93 3.27 4.67 3.15 399.00 263.72 223.89 36.15 520.37 97.71 94.69 99.25 100.00 94.96 45 17 12 17 34 66.12 43.70 37.10 5.99 86.24 0.19 0.93 0.71 0.20 36.17 12.41 65.94 52.62 37.56 31.44 5.26 3.74 6.10 2.85 0.19 0.11 0.33 0.45 0.19 0.93 3.29 1.58 0.19 0.19 0.21 0.19 0.93 0.11 0.19 1.41 0.14 0.33 0.63 0.85 316.55 59.91 254.16 203.18 375.14 2652.1 100. 0 0 100.00 95.69 92.78 93.78 96.89 11 18 15 12 35 21.6 52.46 9.93 42.12 33.67 62.17 43.95

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Appendix 12: Small Benthic Foraminiferal Production Summer 1975 Transects I and II STATION NO. 23 24 25 26 27 28 I 29 30 31 32 33 34 II DEPTH (METERS) 11 18 37 53 90 168 63 19 31 34 37 43 189 59 % LIVING BENTHICS 2.90 11.50 6.00 12.30 4.40 17.20 9.05 12.80 12.60 7.50 14.10 13.90 27.80 14.78 TOTAL LIVING/m2 x 10 5 1.67 0.75 0.95 3.48 1.71 28.35 6.15 13.25 38.10 22.70 3.74 1.99 45.06 20.81 %CARBONATE PRODUCERS 87.67 96.09 95.15 81.97 83.33 94.00 89.70 98.33 89.33 88.67 86.29 85.33 98.33 91.05 CARBONATE PRODUCERS in No. /m 2 X 105 1.46 0.72 0.90 2.85 1.42 26.64 5.67 13.03 34.04 20.12 3.22 1.70 44.31 19.40 Following small species presented in percent of total specimens Ammonia beccarii parkinsoniana 6.00 1.68 1.28 0.00 Ammonia beccarii tepida 13.00 1.12 2.35 0.67 1.33 0.67 0.33 0.50 Amphicoryne intercellularis 0.00 0.00 Amphicoryne sp. 0.00 0.33 0.06 Amphicoryne sublineata 0.33 0.06 0.00 Articulina sulcata 0.34 0.33 0.33 0.17 0.00 Astacolus crepidulus 0.33 0.06 0.33 0.06 Asterigerina carinata 8.94 9.69 0.33 3.16 30.67 2.00 1.33 5.35 6.56 Brizalina albatrossi 0.00 0.00 Brizalina barbata 0.00 0.00 Brizalina fragilis 0.33 0.06 0.33 0.06 Brizalina goessii 0.33 5.67 1.00 1.67 0.28 Brizalina lanceolata 1.67 2.00 0.61 0.67 0.67 0.22 Brizalina lowmani 1.67 0.28 1.00 0.67 2.67 1.33 0.94 Brizalina minima 1.33 0.22 0.33 0.06 Brizalina paula 2.00 0.33 1.33 0.22 Brizalina sp. 0.00 0.00 Brizalina subaenariensis mexicana 0.00 0.33 0.06 Brizalina subspinescens 1.67 0.28 0.67 0.11 Buccella hannai 0.00 0.33 0.06 Bulimina affinis 0.00 1.67 0.28 Bulimina spicata 0.33 0.06 0.00 Buliminella elegantissima 0.33 0.06 0.00

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Appendix 12: (Continued) Cancris oblonga 0.44 0.07 0.67 2.33 1.00 0.67 Cancris sagra 0.44 1.02 0.24 0.33 3.67 0.33 0.72 Carpenteria proteiformis 0.33 0 06 0.00 Cassidulina carinata 1.67 0 28 0.67 0 .11 Cassidulina crassa 0 33 0.06 1.67 0.28 Cassidulina curvata 1.02 5.33 1.33 1.28 0.33 7.33 1.28 Cassidulina subglobosa 2.72 6 67 4 00 2.23 0.67 6.00 1.67 1.39 Chrysalidinella miocenica 0.00 0.33 0 06 Cibicides aff C. floridanus 3 .91 14. 98 11.56 25.33 22.67 13.08 18.00 10.00 4 00 8.36 11.00 13.00 10.73 Cibicides concentricus 0.00 6.33 1.06 Cibicides corpulentus 0.00 1.67 0.28 Cibicides deprimus 3.00 8.94 5 .73 1.70 0.67 1.00 3 .51 2.33 2.67 1.00 0.33 1.06 Cibicides sp. 0.56 0.09 0.00 Cribroelphidium poeyanum 3.91 0.88 0 80 2.67 2 67 2.33 0.67 1.00 1.56 Cyc/ogyra invo/vens 0.00 0.33 0.06 Cyc/ogyra planorbis 0 34 0.06 0 00 Dentalina advena 0.00 0.67 0 .11 Dentalina filiform is 0.00 0 00 Ehrenbergina spinea 1.33 0.22 0.00 Elphidium advenum 0.44 0.07 0.33 0.33 0.11 Elp_hidium discoidale 2.00 1.12 0.44 0.68 0.33 0.76 0.67 1.33 2.33 1.00 0.33 0 .95 Elphidium sp. 4.47 0.34 0.33 0.86 1.67 1.33 0.67 0.61 Eponides antillarum 0.44 0.67 0.18 2.33 3 33 1.00 1.11 Eponides regularis 0.33 0.06 0.00 Eponides turgidus 1.67 0.28 0.33 0.06 Eponides umbonatus 0 00 0.00 Fissurina formosa 0.33 0.06 0.00 Fissurina /ongispina 0.33 0.06 0 00 Fissurina sp. 0.00 0.33 1.33 0.28 Florilus grateloupi 0.56 0.44 0.17 0 33 0.33 0.11 Fursenkoina complanata 0.00 0 00 Fursenkoina compressa 0.88 2.72 0.33 0.66 1.67 2.01 4.33 1.33 Fursenkoina mexicana 0.00 0.67 1.67 2.00 0.72

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Appendix 12: (Continued) Fu r senkoina p ontoni 1.32 0.34 0.28 0.33 0.33 0.67 0.22 Globulina caribaea 0.44 0.07 0.33 0.33 0.11 Guttulina australis 1.12 0.88 0.68 0.45 0.33 0.33 1.67 0.39 Guttulina laevis 0.00 0.33 0.33 0.11 Gyroidina orbicularis 0.00 2.33 0.39 Hanzawaia strattoni 11.67 25.70 10.13 5 78 5.33 1.00 9.94 2.00 10.00 9.33 8 .0 3 2.00 5 .33 6.12 Hoeglundina elegans 0 67 0.11 2.33 0.39 Lenticulina calcar 0.67 0.11 1.67 0.28 Lenticulina gibba 0.33 1.67 0.33 1.33 0 22 Lenticulina orbicularis 1.33 0.22 2.00 0.33 Loxostomum abruptum 0.00 0 .33 0.06 Loxostomum sp. 0.34 0.06 0.33 0.67 0.17 Marginulina glabra 0 00 0.67 0.1 1 Marginulina planata 0 .33 0 06 0.00 Marginulopsis bradyi 0.33 0.06 0.00 Miliolids (abnormal) 0.67 0 .11 0 00 Miliolinella circularis 0 00 0 00 Milioli n el/a obliquinoda 0 33 0.06 0.00 Miliolinel/a subrotunda 0.88 0.33 2.33 0.59 0 .33 1.00 0.33 0.28 Neoconorbina orbicularis 1.68 2.20 19.05 0.33 3.88 0.67 3.67 1.00 1.0 0 6.33 2.11 Nodobacu l ariella cassis 1.00 0.17 0.33 0.33 0.33 0.17 Nonion affinis 0 34 0.06 0.33 0.06 Nonion formosum 0 00 0.33 0.06 Nonio n sp. 0.33 0.33 0.11 0.67 0.67 0.22 Nonionella atlantica 0.44 1.02 0.33 0.3 0 1.00 3.00 1.33 1.34 1.67 0.33 1.45 Nonionella opima 0.00 0.33 0.06 Planorbulina acervalis 0.56 0.68 0.21 0.67 0.33 0.17 P l anorbulina mediterranensis 0.67 0.56 0.44 1.02 0.67 2 00 0.89 0 33 0.33 1.33 0 .33 Planulina ariminensis 5 .33 0.89 9.67 1.61 Planulina exorna 1.00 1.76 3.74 0.67 1.67 1.47 4.33 4.33 3.34 4.00 2 67 Poroeponides latera/is 0.33 1.70 0 33 0.39 0.67 1.33 0.33 Pullenia bulloides 0. 00 0.67 0.11 Pullenia quinqueloba 0.67 0.11 0.00

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........ N 0'\ Pyrgo denticulata Pyrgo depressa Pyrgo elongata Pyrgo nasutus Pyrgo subsphaerica Quinqueloculina bicostata Quinqueloculina bidentata Quinqueloculina bosciana Quinqueloculina compta Quinqueloculina funafutiensis Quinqueloculina horrida Quinqueloculina laevigata Quinqueloculina lamarckiana Quinqueloculina parkeri occidenta/is Quinqueloculina poeyana Quinqueloculina polygona Quinqueloculina seminulum Quinqueloculina sp. Quinqueloculina tenagos Quinqueloculina venusta Quinqueloculina vulgaris Reussella atlantica Rosa/ina bulbosa Rosa/ina columbiensis Rosa/ina concinna Rosa/ina jloridana Rosa/ina jloridensis Rotamorphina laevigata Sagrina pulchel/a primitiva Saracenaria italica Saracenaria latifrons Seabrookia earlandi Sigmavirgu/ina tortuosa 0.33 6.67 6.15 1.00 0.56 0.33 0.67 0.56 0.56 15.67 4.47 12.00 10.61 10.67 3.91 Appendix 12: (Continued) 0.00 0.33 0.06 0.44 0.33 0.13 1.67 2.00 0.61 0.00 0.88 0.15 0.00 1.32 0.33 0.33 0.33 0.00 0.44 0.13 0.67 0.11 0.44 0.07 10.13 2.38 8.33 2.33 6.00 0.00 0.26 0.67 0.17 0.44 0.67 0.33 0.44 0.44 1.02 0.33 0.30 0.09 0.34 0.33 0.11 0.00 0.44 1.70 0.33 2.67 0.86 0.00 12.33 6.12 3.00 0.33 6.99 7.05 3.74 0.33 5.62 0.88 1.02 0.33 2.80 0.34 1.67 0.33 3.00 0.50 0.44 0.07 0.33 0.06 0.00 0.67 0.11 0.00 0.33 0.06 0.00 0.00 1.00 3.33 0.33 0.33 0.83 0.33 0.33 0.33 0.17 0.33 2.34 0.45 0.00 0.67 0.67 0.22 0.67 0.11 0.33 0.06 0.33 0.06 0.00 0.67 6.33 7.67 11.04 2.33 1.33 4.90 0.00 0.33 0.33 0.11 0.33 0.33 0.67 0.33 0.28 0.67 0.11 1.00 0.33 0.22 0.00 0.33 0.33 0.11 0.33 0.06 1.67 3.00 6.35 1.00 1.67 2.28 0.67 0.33 4.00 3.01 0.33 1.39 29.67 17.00 20.67 10.37 13.00 1.33 15.34 2.67 6.67 4.00 2.34 2.33 3.00 1.33 3.33 0.67 2.01 2.00 1.56 1.67 0.67 1.00 0.33 0.61 0.00 0.67 1.67 1.00 0.33 0.61 0.67 0.11 0.00 0.00 0.33 0.06

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Appendix 12: (Continued) Sigmoilina distorta 0.33 0.06 0.33 0.06 Sigmoilina tenuis 0.00 1.00 0.17 Sigmoilopsis schlumbergeri 0.00 0.33 1.33 0.28 Siphonina bradyana 0.33 0.06 2.00 0.33 Siphon ina pulchra 2.67 0.44 0.33 5.00 0.89 Spirillina vivipara 0.00 0.33 0.06 Spirolocu/ina grata 0.00 0.33 1.00 0.33 0 28 Spiroloculina soldanii 0.88 2.00 0.48 0.67 0.11 Spirolocu/ina sp. 0.00 0.33 0.06 Tretomphalus at/anticus? 0.33 0.06 0.00 Tretomphalus bulloides 0.00 0 00 Tretomphalus planus? 0.33 0.06 0.00 Trifarina bella 0.33 0.06 0.33 2.00 0.39 N Trifarina bradyi 0.33 0.67 0.17 0.33 0.06 -.....) Trifarina jamaicensis 0.44 0.34 0.13 0.67 1.00 2.00 0.61 Trilocu/ina linneiana comis 0.00 0.33 0.06 Trilocu/ina sp 0.33 0.06 0.00 Triloculina tricarinata 0.56 0.09 0.33 0.06 Triloculina trigonula 0.33 0.44 0.67 0.24 0.33 0.33 0.11 Triloculina trigonula multistriata 0.00 0.00 Uvigerina bellula 2.00 0.33 0.00 Uvigerina jlintii 2 00 0.33 3.67 0.61 Uvigerina parvula 0.33 1.33 0.28 0 00 Uvigerina peregrina 0.67 0.11 0.33 0.06 Valvulineria minuta 0 00 0 00 Wiesnerella auriculata 0.44 0.34 0.13 0.00 CARBONATE PRODUCERS in No. / m 2 X 105 1.46 0.69 0.86 2.59 1.33 25.23 5.36 13.03 33.53 19.52 3.15 1.63 44.16 1 9 .17 CARBONATE PRODUCTION BY SMALL BENTHICS in g CaCO 3 m -2 yr-1 2.41 1.14 1.43 4.30 2.21 41.81 8.88 21.59 55.57 32.34 5.22 2.70 73.19 31.77

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........ N 00 Appendix 13: Small Benthic Foraminiferal Production Summer 1975 Transect III and Averages STATION NO. 35 36 37 38 39 40 III SUMMER 1975 DEPTH (METERS) 176 43 38 35 29 20 57 60 % LIVING BENTHICS 37.60 2.60 10.60 21.90 12.12 11.98 TOTAL LIVING/m2 x 10 5 29.89 23.59 3.89 10.39 33.31 5.67 17.79 14.92 %CARBONATE PRODUCERS 99.33 89.67 89.67 89.67 91.69 98.33 93.06 91.27 CARBONATE PRODUCERS in No. I m 2 x 10 5 29.69 21.15 3.49 9.32 30.54 5.58 16.63 13.90 Following small species presented in percent of total specimens Ammonia beccarii parkinsoniana 0.33 0.33 0.11 0.46 Ammonia beccarii tepida 1.66 0.28 1.04 Amphicoryne intercellularis 0.33 0.06 0.02 Amphicoryne sp. 0.00 0 .02 Amphicoryne sublineata 0.00 0.02 Articulina sulcata 0.33 0.66 0.17 0.11 Astacolus crepidulus 0.33 0.06 0 .06 Asterigerina carinata 1.00 2.33 32.00 5.89 5.20 Brizalina albatrossi 0.67 0.11 0.04 Brizalina barbata 0.67 0.11 0.04 Brizalina fragilis 0.00 0.04 Brizalina goessii 0.67 0.11 0.46 Brizalina lanceolata 2.33 0.39 0.41 Brizalina lowmani 6.67 0.33 0.33 1.22 0.81 Brizalina minima 0.33 0.06 0.11 Brizalina paula 2.33 0.39 0.31 Brizalina sp. 0.33 0.06 0.02 Brizalina subaenariensis mexicana 0.00 0.02 Brizalina subspinescens 0.00 0.13 Buccella hannai 0.00 0.02 Bulimina ajjinis 0.00 0.09 Bulimina spicata 0.00 0.02 Buliminella elegantissima 0.67 0.11 0.06

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Appendix 13: (Continued) Cancris oblonga 0.00 0.25 Cancris sagra 0.33 0.06 0.34 Carpenteria proteiformis 0.00 0.02 Cassidulina carinata 1.00 0.33 0.33 0.28 0.22 Cassidulina crassa 0.00 0.11 Cassidulina curvata 8.33 0.33 1.44 1.33 Cassidulina subglobosa 7.33 1.33 1.44 1.69 Chrysalidine/la miocenica 0.00 0.02 Cibicides aff. C. jloridanus 11.00 5.67 7.97 13.00 6.27 10.03 Cibicides concentricus 4.00 0.33 0.72 0.59 Cibicides corpulentus 3.00 0.50 0.26 Cibicides deprimus 0.33 2.00 3.32 8.33 2.33 2.30 Cibicides sp. 0.00 0.03 N 10 Cribroelphidium poeyanum 1.00 0.67 0.28 0.88 CyclogJ!!a involvens 0.00 0.02 Cyclogyra planorbis 0.00 0.02 Dentalina advena 0.00 0.04 Dentalina filiformis 0.33 0.06 0.02 Ehrenbergina spinea 0.00 0.07 Elphidium advenum 0.00 0.06 Elphidium discoidale 3.67 1.66 0.89 0.86 Elphidium sp. 0.33 0.06 0.51 Eponides antil/arum 1.33 2.33 0.61 0.64 Eponides regularis 0.00 0.02 Eponides turgidus 0.00 0.11 Eponides umbonatus 0.33 0.06 0.02 Fissurina formosa 0.00 0.02 Fissurina longispina 0.00 0.02 Fissurina sp 1.00 0.33 0.33 0.28 0.19 Florilus grateloupi 0.00 0.09 Fursenkoina complanata 0.33 0.06 0.02 Fursenkoina compressa 0.67 0.33 0.17 0.72 Fursenkoina mexicana 0.33 0.33 0.33 0.17 0.30

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_. w 0 Fursenkoina pontoni Globulina caribaea Guttulina australis Guttulina laevis Gyroidina orbicularis Han zawai a strattoni Hoeglundina elegans Lenticulina calcar Lenticulina gibba Lenticulina orbicularis Loxostomum abruptum Loxostomum sp. Marginu/ina glabra Marginulina planata Marginulopsis bradyi Miliolids (abnormal) Miliolinella circularis Miliolinella obliquinoda Miliolinella subrotunda Neoconorbina orbicularis Nodobaculariella cassis Nonion affinis Nonion formosum Nonion sp. Nonionel/a atlantica Nonionel/a opima Planorbulina acervalis Planorbulina mediterranensis Planulina ariminensis Planulina exorna Poro eponides latera/is Pullenia bulloides Pullenia quinqueloba Appendix 13: (Continued) 0.33 0.33 0.67 0.33 1.00 0.33 4.33 8.00 11.30 1.67 2.33 2 00 2.33 0.33 0.33 1.00 1.00 4.67 2.33 0.33 0.67 0.33 3.00 3.33 1.00 7.33 10.33 1.00 0.33 0.06 0.18 0.06 0.08 0.11 0.32 0 22 0.11 0.06 0.15 3.67 4.55 6.87 0 .28 0.26 0.39 0.26 0.33 0.30 0.39 0.31 0.00 0.02 0.06 0.09 0.00 0 04 0.00 0.02 0 00 0.02 0.00 0 04 0.06 0.02 0.00 0 02 0.33 0.39 0.42 2.33 1.55 2.51 0.06 0.13 0.00 0.04 0.11 0.06 0.06 0.13 3.00 1.00 0.91 0.00 0.02 0.00 0.12 0.72 0.65 1.22 1.24 1.89 2.01 0.00 0.24 0.00 0 04 0.06 0 06

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Appendix 13: (Continued) Pyrgo denticulata 0.00 0 .02 Pyrgo depressa 0 33 0 06 0.04 Pyrgo elongata 0.67 0.11 0.08 Pyrgo nasutus 0 .33 0.66 0.17 0.54 Pyrgo subsphaerica 1.00 0.17 0.11 Quinqueloculina b ic ostata 1.67 3.99 0.94 0.51 Quinqueloculina bidentata 0.33 0.06 0.02 Quinqueloculina bosciana 1.00 0.33 0.33 0.33 0 33 0.30 Quinqueloculina compta 0.00 0.04 Quinqueloculina funafutiensis 0.33 1.33 0 .28 0 .15 Quinqueloculina horrida 0 00 0.06 Quinqueloculina la e vigata 0 33 0.06 0.04 Quinqueloculina lamarckiana 0.33 6.33 12.62 3.22 4.70 Quinqueloculina parkeri occidentalis 0 33 0.06 0.02 Quinqueloculina poeyana 0.67 0.11 0.16 Quinqueloculina polygona 0.66 0.11 0.19 Quinqueloculina seminulum 0 00 0.19 Quinqueloculina sp 0.00 0.17 Quinqueloculina tenagos 0 33 0 06 0 05 Quinqueloculina venusta 0 00 0 07 Quinque/oculina vulgaris 0.00 0.02 Reussel/a atlantica 0.33 2.67 1.99 0 .83 1.32 Rosa/ina bulbosa 0.33 0 66 0.17 0.52 Rosa/ina columbiensis 16.00 18. 27 17.67 8.66 10.33 Rosa/ina concinna 4.00 1.66 10.67 2.72 3.78 Rosa/ina jloridana 2.33 2 66 1.00 1.00 1.79 Rosa/ina jloridensis 0 33 0 06 0.33 Rotamorphina laevigata 0.00 0.17 Sagrina pulchel/a primitiva 0 .33 1.00 0.22 0.30 Saracenaria italica 1.00 0 .17 0.11 Saracenaria latifj(Jns 0.33 0.06 0 02 Seabrookia earlandi 0.00 0.04 Sigmavirgulina tortuosa 0.00 0.02

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........ w N Sigmoilina distorta Sigmoilina tenuis Sigmoilopsis schlumbergeri Siphonina bradyana Siphon ina pulchra Spirillina vivipara Spirolocu/ina grata Spiro/oculina soldanii Spiroloculina sp Tretomphalus at/anticus? Tretomphalus bulloides Tretomphalus planus? Trifarina bella Trifarina bradyi Trifarina jamaicensis Triloculina linneiana comis Triloculina sp. Triloculina tricarinata Triloculina trigonula Triloculina trigonula multistriata Uvigerina bellula Uvigerina flintii Uvigerina parvula Uvigerina peregrina Valvu/ineria minuta Wiesnerella auriculata CARBONATE PRODUCERS in No. I m 2 x I 0 5 CARBONATE PRODUCTION BY SMALL BENTHICS ingCaC03 m -2 -1 yr Appendix 13: (Continued) 0.33 10.33 0.33 0.33 0.33 0.33 0.33 0.33 0.67 0.33 0.33 0.66 6.33 1.00 0.67 29.59 20.60 3.40 9.08 29.65 49.04 34.14 5.63 15.04 49.14 0.00 0.04 0.06 0 .07 0.00 0.0 9 0.00 0.13 1.72 1.02 0.00 0 0 2 0.06 0.11 0.00 0.2 0 0.00 0.02 0.06 0.04 0.06 0.02 0.06 0.04 0.11 0.19 0.00 0.07 0.11 0.28 0.00 0.02 0.00 0.02 0.06 0.07 0.33 0.22 0.19 0.33 0.06 0.02 0.00 0.11 1.06 0.67 0.00 0.09 0.17 0.11 0.11 0.04 0.00 0.04 5.43 16.29 13.61 8.99 27.00 22.55

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..... w w Appendix 14: Small Benthic Foraminiferal Production Falll975 Transects I, II, III and Averages STATION NO. 23 24 25 26 27 28 29 30 3I 32 33 34 35 36 37 38 DEPTH (METERS) II I8 37 53 90 I68 I9 3I 34 37 43 I89 I76 43 29 35 % LIVING BENTHICS I5.I 13.4 I0.8 6.5 I1.6 22.7 I2.I I3.3 I9.7 I7.I Il.7 20.6 39.3 I4.I 21.7 I0.5 TOTAL LIVING/m2 x I05 30.9 I7.7 I3.0 2.2 Il.8 26.0 I5.I 43.7 70.6 30.2 Il.7 I0.2 63.6 5.4 54.4 1.6 %CARBONATE PRODUCER 87.7 96.I 95.2 82.0 83.3 94.0 98.3 89.3 88.7 86.3 85.3 98.3 99.3 91.5 91.5 89.7 CARBONATE PRODUCERS 27.I I7.0 12.4 1.8 9.9 24.5 I4.8 39.I 62.6 26.0 IO.O IO.I 63.2 5.0 49.7 I.4 in No. 1m2 x 105 Ammonia beccarii parkinsonian 6.0 1.7 0.3 Ammonia beccarii tepida 13.0 1.1 0.7 1.3 0.7 0.3 Amphicoryne intercel/ularis 0.3 Amphicoryne sp 0.3 Amphicoryne sublineata 0.3 Articulina su/cata 0.3 0.3 0.3 0.3 Astacolus crepidulus 0.3 0.3 0.3 Asterigerina carinata 8.9 9.7 0.3 30.7 2.0 1.3 5.4 1.0 Brizalina a/batrossi 0.7 Brizalina barbata 0.7 Brizalina fragilis 0.3 0.3 Brizalina goessii 0.3 5.7 1.7 0.7 Brizalina /anceolata 1.7 2.0 0.7 0.7 2.3 Brizalina lowmani 1.7 1.0 0.7 2.7 1.3 6.7 0.3 Brizalina minima 1.3 0.3 0.3 Brizalina y_aula 2.0 1.3 2.3 Brizalina sp. 0.3 Brizalina subaenariensis mexicana 0.3 Brizalina subspinescens 1.7 0.7 Buccel/a hannai 0.3 Bulimina affinis 1.7 Bulimina spicata 0.3 Buliminella elegantissima 0.3 0.7 Cancris oblonga 0.4 0.7 2.3 1.0 Cancris sagra 0.4 1.0 0.3 3.7 0.3 0.3 39 40 29 20 59 Il.3 49.6 I7.8 4.7 6.4 23.3 91.7 98.3 91.5 4.3 6.3 21.4 0.3 1.7 0.7 2.3 32.0 0.3

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Appendix 14: (Continued) Carpenteria proteiformis 0.3 Cassidulina carinata 1.7 0.7 1.0 0.3 0 3 Cassidulina crassa 0.3 1.7 Cassidulina curvata 1.0 5.3 1.3 0.3 7.3 8 3 0.3 Cassidulina subglobosa 2.7 6.7 4.0 0.7 6.0 1.7 7 3 1.3 Chrysalidinella miocenica 0 3 Cibicides aff. C. floridanus 3.9 15. 0 11.6 25.3 22.7 18.0 10. 0 4.0 8.4 11.0 13. 0 11.0 5 7 8.0 13.0 Cibicides concentricus 6.3 4 0 0.3 Cibicides corpulentus 1.7 3.0 Cibicides deprimus 3.0 8.9 5.7 1.7 0.7 1.0 2.3 2.7 1.0 0.3 0.3 2.0 3.3 8.3 Cibicides sp. 0.6 Cribroelphidium poeyanum 3.9 0.9 2.7 2 7 2.3 0.7 1.0 1.0 0.7 Cyclogyra involvens 0.3 Cyclogyra planorbis 0.3 Dentalina advena 0.7 Dentalina filiformis 0 3 Ehrenbergina spinea 1.3 Elphidium advenum 0.4 0.3 0.3 Elphidium discoida/e 2.0 1.1 0.4 0 7 0.3 0.7 1.3 2.3 1.0 0.3 3.7 1.7 Elp_hidium sp. 4.5 0.3 0.3 1.7 1.3 0.7 0.3 Eponides antillarum 0.4 0.7 2.3 3.3 1.0 1.3 2.3 Eponides regu/aris 0.3 Eponides turgidus 1.7 0.3 Eponides umbonatus 0 3 Fissurina formosa 0.3 Fissurina longispina 0.3 Fissurina sp. 0.3 1.3 1.0 0 3 0.3 Florilus grateloupi 0.6 0.4 0.3 0.3 Fursenkoina comp/anata 0.3 Fursenkoina compressa 0.9 2.7 0.3 1.7 2.0 4.3 0.7 0.3 Fursenkoina mexicana 0.7 1.7 2.0 0.3 0.3 0.3 Fursenkoina pontoni 1.3 0.3 0.3 0.3 0.7 0.3 Globulina caribaea 0.4 0.3 0.3 0.3

PAGE 145

Appendix 14: (Continued) Guttu/ina australis 1.1 0.9 0.7 0.3 0.3 1.7 0.7 Guttulina laevis 0.3 0.3 0.3 1.0 G y roidina orbicularis 2.3 0.3 Hanzawaia strattoni 11.7 25.7 10.1 5.8 5.3 1.0 2.0 10.0 9.3 8.0 2.0 5.3 4.3 8.0 11.3 3.7 Hoeglundina elegans 0.7 2.3 1.7 Lenticulina calcar 0.7 1.7 2.3 Lenticulina gibba 0.3 1.7 1.3 2.0 Lenticulina orbicularis 1.3 2.0 2.3 Loxostomum abruptum 0.3 Loxostomum s p 0.3 0.3 0.7 0.3 Marginulina glabra 0.7 Marginulina planata 0.3 Marginulopsis bradyi 0.3 Milio/ids (abnormal) 0.7 Miliolinella circularis 0.3 -w VI Miliolinel/a obliquinoda 0.3 Miliolinella subrotunda 0.9 0.3 2.3 0.3 1.0 0.3 1.0 1.0 0.3 Neoconorbina orbicularis 1.7 2.2 19.0 0.3 0.7 3.7 1.0 1.0 6.3 4.7 2.3 2.3 Nodobaculariella cassis 1.0 0 3 0.3 0.3 0.3 Nonion a./finis 0.3 0.3 Non ion formosum 0.3 0 7 Nonion sp. 0.3 0.3 0.7 0.7 0.3 Nonione/la atlantica 0.4 1.0 0.3 1.0 3.0 1.3 1.3 1.7 0.3 3.0 3.0 Nonionel/a opima 0.3 Planorbu/ina acerva/is 0.6 0 7 0.7 0.3 Planorbulina mediterranensis 0.7 0.6 0.4 1.0 0 7 2.0 0.3 0 3 1.3 3.3 1.0 Planulina ariminensis 5.3 9.7 7.3 Planu/ina exorna 1.0 1.8 3.7 0 7 1.7 4.3 4.3 3.3 4.0 10.3 1.0 Poroeponides latera/is 0.3 1.7 0.3 0.7 1.3 Pullenia bulloides 0.7 Pullenia quinqueloba 0.7 0.3 IPyrgo denticulata 0.3 IPvrR"O depressa 0.3 0.3

PAGE 146

Appendix 14: (Continued) Pyrgo elongata 0.4 0.3 0.7 IPyrgo nasutus 1.7 2.0 1.0 3.3 0.3 0.3 0.3 0.7 Pyrgo subsphaerica 0.3 0.3 0.3 1.0 Quinqueloculina bicostata 0.9 0.3 2.3 1.7 4.0 Quinqueloculina bidentata 0.3 Quinqueloculina bosciana 1.3 0.3 0.3 0.7 0.7 1.0 0.3 0.3 0.3 Quinqueloculina compta 0.7 Quinqueloculina funafutiensis 0.3 0.4 0.3 0.3 1.3 Quinqueloculina horrida 0.7 0.3 Quinqueloculina laevigata 0.4 0.3 Quinqueloculina lamarckiana 6.7 6.1 10.1 2.4 8.3 2.3 0.7 6.3 7.7 11.0 2.3 1.3 0.3 6.3 12.6 Quinqueloculina parkeri occidentalis 0.3 Quinqueloculina poeyana 1.0 0.6 0.3 0.3 0.7 Quinqueloculina polygona 0.3 0.7 0.3 0.3 0.7 0.3 0.7 Quinqueloculina seminulum 0.7 0.6 0.4 0.7 0.3 0.7 Quinqueloculina sp. 0.4 1.0 0.3 1.0 0.3 Quinqueloculina tenagos 0.6 0.3 Quinqueloculina venusta 0.3 0.3 0.3 0.3 Quinqueloculina vulgari s 0.3 Reussella atlantica 0.4 1.7 0.3 2.7 1.7 3.0 6.4 1.0 1.7 0.3 2.7 2.0 Rosa/ina bulbosa 0.7 0.3 4.0 3.0 0.3 0.3 0.7 Rosa/ina columbiensis 15.7 4.5 12.3 6.1 3.0 0.3 29.7 17.0 20.7 10.4 13.0 1.3 16.0 18.3 17.7 Rosa/ina concinna 12.0 10.6 7.0 3.7 0.3 2.7 6.7 4.0 2.3 2.3 4.0 1.7 10.7 Rosa/ina floridana 10.7 3.9 0.9 1.0 0.3 1.3 3.3 0.7 2.0 2.0 2.3 2.7 1.0 Rosa/ina florid ens is 0.3 1.7 1.7 0.7 1.0 0.3 0.3 Rotamorphina laevigata 3.0 Sagrina pulchella primitiva 0.4 0.7 1.7 1.0 0.3 0.3 1.0 Saracenaria italica 0.3 0.7 1.0 Saracenaria latifrons 0.3 Seabrookia earlandi 0.7 Sigmavirgulina tortuosa 0.3 Sigmoilina distorta 0.3 0.3 Sigmoilina tenuis 1.0 0.3

PAGE 147

Appendix 14: (Continued) Sigmoilopsis schlumbergeri 0.3 1.3 Siphonina bradyana 0.3 2.0 Siphon ina pulchra 2.7 0.3 5.0 10.3 Spirillina vivipara 0.3 Spiroloculina grata 0.3 1.0 0.3 0.3 Spiroloculina soldanii 0.9 2.0 0.7 Spiroloculina sp. 0.3 Tretomphalus at/anticus? 0.3 0.3 Tretomphalus bulloides 0.3 Tretomphalus planus? 0.3 0.3 Trifarina bella 0.3 0.3 2.0 0.3 0.3 Trifarina bradyi 0.3 0.7 0.3 Trifarina jamaicensis 0.4 0.3 0.7 1.0 2.0 0.7 Triloculina linneiana comis 0.3 Triloculina sp. 0.3 Triloculina tricarinata 0.6 0.3 0.3 Triloculina trigonula 0.3 0.4 0.7 0.3 0.3 0.3 0.7 0.3 Triloculina trigonula multistriata 0.3 Uvigerina bellula 2.0 Uvigerina flintii 2.0 3.7 6.3 Uvigerina parvula 0.3 1.3 Uvigerina peregrina 0.7 0.3 1.0 Valvulineria minuta 0.7 Wiesnerella auriculata 0.4 0.3 SMALL CARBONATE PROD 27.0 16.3 11.9 1.6 9.2 23.2 14.8 38.5 60.7 25.4 9.6 10.0 63.0 4.8 48.2 1.4 4.1 6.1 20.9 CARBONATE PRODUCTION ing CaC03 m 2 -1 44.8 27.0 19.7 2.7 15.3 38.4 24.6 63.8 yr 100.6 42.2 15.9 16.6 104.4 8.0 79.9 2.3 6.9 10.1 34.6

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...... w 00 Appendix 15: Small Benthic Foraminiferal Production Winter 1976 Transects I, II, III and Averages STATION NO. 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 DEPTH (METERS) 11 18 37 53 90 168 19 31 34 37 43 189 176 43 29 35 29 % LIVING BENTHICS 15.4 17.8 6.7 18.8 6.9 9.7 9.5 9.7 9.1 16.0 13.4 24.3 27.3 30.8 19.6 16.3 8.2 TOTAL LIVING/m2 x 10 5 4.3 11.1 11.4 15.8 2.2 6.9 7.0 15.6 14.8 9.0 2.4 6.3 41.6 5.7 4.2 0.7 16.5 %CARBONATE PRODUCER 91.5 91.5 91.5 91.5 91.5 91.5 91.5 91.5 91.5 91.5 91.5 91.5 91.5 91.5 91.5 91.5 91.5 CARBONATE PRODUCERS 3.9 10.2 10.4 14.5 2.0 6.3 6.4 14.3 13.5 8.3 2.2 5.7 38.1 5.2 3.9 0.7 15.1 in No. I m 2 x 10 5 Ammonia beccarii parkinsonian 6.0 1.7 0.3 Ammonia beccarii tepida 13.0 1.1 0.7 1.3 0.7 0.3 1.7 Amphicoryne intercellularis 0.3 Amphicoryne sp. 0.3 Amphicoryne sublineata 0.3 Articulina sulcata 0.3 0.3 0.3 0.3 0.7 Astacolus crepidulus 0.3 0.3 0.3 Asterigerina carinata 8.9 9.7 0.3 30.7 2.0 1.3 5.4 1.0 2.3 Brizalina albatrossi 0.7 Brizalina barbata 0.7 Brizalina fragilis 0.3 0.3 Brizalina goessii 0.3 5.7 1.7 0.7 Brizalina lanceolata 1.7 2.0 0.7 0.7 2.3 Brizalina lowmani 1.7 1.0 0.7 2.7 1.3 6.7 0.3 Brizalina minima 1.3 0.3 0.3 Brizalina paula 2.0 1.3 2.3 Brizalina sp. 0.3 Brizalina subaenariensis mexicana 0.3 Brizalina subspinescens 1.7 0.7 Buccella hannai 0.3 Bulimina affinis 1.7 Bulimina spicata 0.3 Buliminella elegantissima 0.3 0.7 Cancris oblonga 0.4 0.7 2.3 1.0 Cancris sagra 0.4 1.0 0.3 3.7 0.3 0.3 40 20 59 40.1 16.6 1.4 9.8 91.5 91.5 1.3 9.0 0.3 32.0 0.3

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Appendix 15: (Continued) Carpenteria proteiformis 0.3 Cassidulina carinata 1.7 0.7 1.0 0.3 0.3 Cassidulina crassa 0.3 1.7 Cassidulina curvata 1.0 5.3 1.3 0.3 7.3 8.3 0.3 Cassidulina 2.7 6.7 4.0 0.7 6.0 1.7 7.3 1.3 Chrysalidinella miocenica 0.3 Cibicides aff. C. jloridanus 3.9 15.0 11.6 25.3 22.7 18.0 10.0 4.0 8.4 11.0 13.0 11.0 5.7 8.0 13.0 Cibicides concentricus 6.3 4.0 0.3 Cibicides corpulentus 1.7 3.0 Cibicides deprimus 3.0 8.9 5.7 1.7 0.7 1.0 2.3 2.7 1.0 0.3 0.3 2.0 3.3 8.3 Cibicides sp. 0.6 Cribroe/phidium poeyanum 3.9 0.9 2.7 2.7 2.3 0.7 1.0 1.0 0.7 Cyclogyra involvens 0.3 Cyclogyra planorbis 0.3 Dentalina advena 0.7 Dentalina filiformis 0.3 Ehrenbergina spinea 1.3 E/phidium advenum 0.4 0.3 0.3 Elphidium discoidale 2.0 1.1 0.4 0.7 0.3 0.7 1.3 2.3 1.0 0.3 3.7 1.7 Elphidium sp. 4.5 0.3 0.3 1.7 1.3 0.7 0.3 Eponides antillarum 0.4 0.7 2.3 3.3 1.0 1.3 2.3 Eponides regularis 0.3 Eponides turgidus 1.7 0.3 Eponides umbonatus 0.3 Fissurina formosa 0.3 Fissurina 0.3 Fissurina sp. 0.3 1.3 1.0 0.3 0.3 Florilus grateloupi 0.6 0.4 0.3 0.3 Fursenkoina complanata 0.3 Fursenkoina compressa 0.9 2.7 0.3 1.7 2.0 4.3 0.7 0.3 Fursenkoina mexicana 0.7 1.7 2.0 0.3 0.3 0.3 Fursenkoina pontoni 1.3 0.3 0.3 0.3 0.7 0.3 Globulina caribaea 0.4 0.3 0.3 0.3

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Appendix 15: (Continued) Guttulina australis 1.1 0.9 0.7 0.3 0.3 1.7 0.7 Guttulina laevis 0.3 0.3 0.3 1.0 Gyroidina orbicularis 2.3 0.3 Hanzawaia strattoni 11.7 25.7 10.1 5.8 5.3 1.0 2.0 10.0 9.3 8.0 2.0 5.3 4.3 8.0 11.3 3.7 Hoeglundina elegans 0.7 2.3 1.7 Lenticulina calcar 0.7 1.7 2.3 Lenticulina gibba 0.3 1.7 1.3 2.0 Lenticulina orbicularis 1.3 2.0 2.3 Loxostomum abruptum 0.3 Loxostomum sp 0.3 0.3 0.7 0.3 Marginulina glabra 0.7 Marginulina planata 0.3 Marginulopsis bradyi 0.3 Miliolids (abnormal) 0.7 Miliolinella circularis 0.3 Miliolinella obliquinoda 0.3 Miliolinella subrotunda 0.9 0.3 2.3 0.3 1.0 0.3 1.0 1.0 0.3 Neoconorbina orbicularis 1.7 2.2 19.0 0.3 0.7 3.7 1.0 1.0 6.3 4.7 2.3 2.3 Nodobaculariella cassis 1.0 0.3 0.3 0.3 0.3 Nonion affinis 0.3 0.3 Nonion formosum 0.3 0.7 Nonionsp 0.3 0.3 0.7 0.7 0.3 Nonionella atlantica 0.4 1.0 0.3 1.0 3.0 1.3 1.3 1.7 0.3 3.0 3.0 Nonionella opima 0.3 Planorbulina acervalis 0.6 0.7 0.7 0.3 Planorbulina mediterranensis 0.7 0.6 0.4 1.0 0.7 2.0 0.3 0.3 1.3 3.3 1.0 Planulina ariminensis 5.3 9.7 7.3 Planulina exorna 1.0 1.8 3.7 0.7 1.7 4.3 4.3 3.3 4.0 10.3 1.0 Poroeponides latera/is 0.3 1.7 0.3 0.7 1.3 Pullenia bulloides 0.7 Pullenia quinqueloba 0.7 0.3 Pyrgo denticulata 0.3 IPyrgo depressa 0.3 0.3

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Appendix 15: (Continued) Pyrgo elongata 0.4 0.3 0.7 Pyrgo nasutus 1.7 2.0 1.0 3.3 0.3 0.3 0.3 0.7 Pyrgo subsphaerica 0.3 0.3 0.3 1.0 Quinqueloculina bicostata 0.9 0.3 2.3 1.7 4.0 Quinqueloculina bidentata 0.3 Quinquelocu/ina bosciana 1.3 0.3 0.3 0.7 0.7 1.0 0.3 0.3 0.3 Quinquelocu/ina compta 0.7 Quinquelocu/ina funafutiensis 0.3 0.4 0.3 0.3 1.3 Quinqueloculina horrida 0.7 0.3 Quinqueloculina laevigata 0.4 0.3 Quinqueloculina lamarckiana 6.7 6.1 10.1 2.4 8.3 2.3 0.7 6.3 7.7 11.0 2.3 1.3 0.3 6.3 12.6 Quinqueloculina parkeri occidentalis 0.3 Quinqueloculina poeyana 1.0 0.6 0.3 0.3 0.7 Quinqueloculina polygona 0.3 0.7 0.3 0.3 0.7 0.3 0.7 Quinqueloculina seminulum 0.7 0.6 0.4 0.7 0.3 0.7 Quinqueloculina sp. 0.4 1.0 0.3 1.0 0.3 Quinquelocu/ina tenagos 0.6 0.3 Quinqueloculina venusta 0.3 0.3 0.3 0.3 Quinqueloculina vulgaris 0.3 Reussella atlantica 0.4 1.7 0.3 2.7 1.7 3.0 6.4 1.0 1.7 0.3 2.7 2.0 Rosa/ina bulbosa 0.7 0.3 4.0 3.0 0.3 0.3 0.7 Rosa/ina columbiensis 15.7 4.5 12.3 6.1 3.0 0.3 29.7 17.0 20.7 10.4 13.0 1.3 16.0 18.3 17.7 Rosa/ina concinna 12.0 10.6 7.0 3.7 0.3 2.7 6.7 4.0 2.3 2.3 4.0 1.7 10.7 Rosa/ina floridana 10.7 3.9 0.9 1.0 0.3 1.3 3.3 0.7 2.0 2.0 2.3 2.7 1.0 Rosa/ina floridensis 0.3 1.7 1.7 0.7 1.0 0.3 0.3 Rotamorphina laevigata 3.0 Sagrina pulchel/a primitiva 0.4 0.7 1.7 1.0 0.3 0.3 1.0 Saracenaria italica 0.3 0.7 1.0 Saracenaria latifrons 0.3 Seabrookia earlandi 0.7 Sigmavir_gulina tortuosa 0.3 Sigmoilina distorta 0.3 0.3 Sigmoilina tenuis 1.0 0.3

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Appendix 15: (Continued) Sigmoilopsis schlumbergeri 0 3 1.3 Siphonina bradyana 0.3 2.0 Siphonina pulchra 2.7 0.3 5.0 10.3 Spirillina vivipara 0 3 Sp_jrolocu/ina grata 0 3 1.0 0.3 0.3 Spirolocu/ina soldanii 0.9 2 0 0.7 Spiroloculina sp 0 3 Tretomphalus at/anticus? 0 3 0.3 Tretomphalus bulloides 0.3 Tretomphalus planus? 0.3 0.3 Trifarina bella 0.3 0.3 2.0 0 3 0.3 Trifarina bradyi 0.3 0 7 0.3 Trifarina jamaicensis 0.4 0.3 0.7 1.0 2.0 0 7 Triloculina linneiana comis 0 3 .J::. Triloculina sp 0.3 N Triloculina tricarinata 0.6 0.3 0 3 Triloculina trigonula 0.3 0.4 0.7 0.3 0.3 0.3 0.7 0.3 Triloculina trigonula multistriata 0 3 Uvigerina bellula 2.0 Uvigerina flintii 2.0 3.7 6.3 Uvigerina parvula 0.3 1.3 Uvigerina peregrina 0.7 0.3 1.0 Valvulineria minuta 0.7 Wiesnerella auriculata 0.4 0.3 CARBONATE PRODUCERS in No. 1m2 x 105 3.8 9.9 10.1 14. 0 2 0 6.2 6 2 13. 9 13. 1 8 0 2.1 5.6 36.9 5 1 3.7 0 6 14. 6 1.3 8.7 CARBONATE PRODUCTION BY SMALL BENTHICS Winter 1976 ingCaC03 m -2 yr 1 6.3 16.4 16.8 23.3 3.3 10.2 10.2 23.0 21.7 13.3 3.5 9.2 61.2 8.4 6.2 1.1 24.3 2.1 14.5

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Appendix 16: Foraminiferal Production Rates by Season 1975-1976 Large Small Station 1975 1976 1975 1976 Total Total Total Total Summer Fall Winter Avg. Summer Fall Winter Avg Summer Fall Winter 23 0.17 3.09 0.43 1.23 2.41 44.78 6.30 17.83 2.58 47.87 6.73 19.06 24 0.80 18.94 11.93 10.56 1.14 26.98 16.40 14.84 1.94 45.92 28.33 25.40 25 0.45 6.20 5.43 4.03 1.43 19.70 16.80 12.64 1.88 25.90 22.23 16.67 26 5.73 3.57 26.03 11.78 4.30 2.68 23.30 10.09 10.03 6.25 49.33 21.87 27 2.97 20.61 3.89 9.16 2.21 15.31 3.30 6.94 5 .18 35.92 7.19 16.10 28 46.11 42.35 11.28 33.25 41.81 38.40 10.20 30.14 87 92 80.75 21.48 63.38 9.37 15.79 9.83 11.67 8.88 24.64 12.72 15.41 18.26 40.44 22.55 27.08 29 0.00 0.00 0.00 0.00 21.59 24.61 10.20 18.80 21.59 24.61 10.20 18.80 30 6 10 7.00 2.50 5.20 55.57 63.77 23.00 47.45 61.67 70.77 25.50 52.65 31 7.26 22.58 4.73 11.52 32.34 100.57 21.70 51.54 39.60 123.15 26.43 63 06 32 1.17 9.49 2.84 4.50 5.22 42.15 13.30 20.22 6.39 51.64 16.14 24.72 33 2.25 13.29 2.67 6.07 2.70 15.93 3.50 7.38 4.95 29.22 6.17 13.45 34 1.80 0.41 0.25 0.82 73.19 16.63 9.20 33.01 74.99 17.04 9.45 33.83 3.10 8.80 2.17 4.69 31.77 43.94 13.48 29.73 34.87 52.74 15.65 34.42 35 3.39 7.21 4.72 5.11 49.04 104.39 61.20 71.54 52.43 111.60 65.92 76.65 36 13.52 1.27 0.65 5 .15 34.14 7.86 8.40 16.80 47.66 9.13 9.05 21.95 37 2.23 12.68 0.48 5.13 5.63 79.87 6.20 30.57 7.86 92.55 6.68 35 70 38 5.96 0.92 0.41 2.43 15.04 2.33 1.10 6.16 21.00 3.25 1.51 8.59 39 14.61 2.04 7.24 7.96 49.14 6.87 24.30 26.77 63.75 8.91 31.54 34.73 40 3.52 3.97 0.90 2.80 8.99 10.14 2.10 7.08 12.51 14.11 3.00 9.87 7.21 4.68 2.40 4.76 27.00 35.24 17.22 26.49 34.20 39.93 19.62 31.25 Avg. 6.56 9 .76 4.80 7.04 22.55 34.61 14.47 23.88 29.11 44.37 19.27 30.91

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Avg. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 37 45 44 53 37 36 40 42 48 54 37 34 44 38 38 43 31 33 34 30 30 42 11 18 37 53 90 168 19 31 34 37 43 189 176 29 38 35 29 20 48 Appendix 17: Foraminiferal Production Rate Summary Large Foraminifera 1 7 19 12 9 8 8 9 12 17 11 8 3 5 6 12 18 5 0 0 12 17 16 15 2 6 5 11 1 8 0 0 5 8 0 0 0 0 4 12 7 14 6 10 0 4 4 12 4 3 9 9 6 4 5 4 0 0 1 5 3 5 2 7 4 3 0 2 0 2 3 7 3 7 3 7 3 9 3 17 5 7 3 7 36.1 12.2 15. 1 15.4 7.8 21.7 13.3 47.6 0 0 18. 6 7 .9 21.5 17.7 2 0 0.0 37.4 0 0 0.0 23. 2 86.4 29.9 1.2 10.6 4.0 11. 8 9.2 33.3 0.0 5.2 11.5 4.5 6 1 0.8 5.1 5 2 5.1 2.4 8.0 2.8 13. 6 Small Foraminifera 99 81 91 92 88 89 97 94 82 100 88 84 98 95 99 100 95 100 100 96 93 94 100 96 96 91 94 95 100 99 97 98 96 100 100 97 97 97 97 97 95 144 93 47 5 88 10.4 92 6.6 91 22.4 83 10.8 92 4.3 95 31.2 88 11.0 95 10.5 100 86 1 83 13.0 85 3.6 94 66. 1 89 43 .7 92 37 1 100 6.0 92 86.2 100 52.5 100 9.9 88 42 1 86 33.7 90 62 2 96 17.8 88 14 8 97 12.6 91 10.1 96 6.9 96 30.1 100 18.8 95 47 5 95 51.5 93 20.2 98 7.4 98 33.0 98 71.5 93 16 8 93 30.6 93 6.2 91 26.8 83 7 1 93 28 2 Total Foraminifera 51.2 46.5 18.9 37.4 26.2 12. 1 52.9 24 3 58.1 86.1 31.6 11.5 87.6 61.4 39.1 6 0 123.6 52 5 9 9 65.3 120 0 92 0 19.1 25.4 16.7 21.9 16.1 63.4 18.8 52.7 63. 1 24.7 13.5 33 8 76.7 22 0 35.7 8 6 34 .7 9.9 41.8 7.3 77.6 64.9 40.3 58.9 64.4 40.9 54.8 82 0 0.0 58.8 68.7 24.5 28.9 5.2 0.0 30.2 0.0 0.0 35 5 72.0 32.4 6.5 41.6 24.2 53.9 56.9 52.5 0.0 9.9 18. 3 18.2 45.1 2.4 6.7 23.5 14.4 28.3 22.9 28.3 32.5 92.7 22.4 35.1 59.7 41.1 35 6 59.1 45 2 18. 0 100.0 41.2 31.3 75. 5 71.1 94.8 100.0 69.8 100.0 100.0 64.5 28.0 67.6 93. 5 58.4 75.8 46.1 43.1 47.5 100.0 90.1 81.7 81.8 54.9 97.6 93.3 76.5 85.6 71.7 77.1 71.7 67 5

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Appendix 18: Micromolluscan Production and Species List 1974 Transect II STATION NO. 1 2 3 4 5 6 7 8 9 10 11 12 STA. DEPTH (METERS) 37 45 44 53 37 36 40 42 48 54 37 34 NO. SPECIMENS x 104 /m2 23.5 9.1 25.0 19.5 33.0 35.4 38.5 20.1 17.4 11.6 33.7 28.3 24.6 NO. LIVING SPEC x 102 /m2 14.1 5.4 15.0 11. 7 19.8 21.2 23.1 12.1 10.4 7.0 20.2 17.0 14.7 CALCIMASS in g CaC03 m-2 5.1 2.0 5.4 4.2 7.2 7.7 8.4 4.4 3.8 2.5 7.3 6.2 5.3 CaC03 PRODUCTION RATE in g CaC03 m-2 yr1 51.08 19.69 54.47 42.47 71.70 76.93 83.70 43.70 37.85 25.23 73.24 61.54 53.47 BIVALVES 0 Abra lioica 1 0 Arcopsis adamsi 1 1 3 0 Astarte nana 1 1 0 Cardiomya ornatissima 1 0 Cardiomya perrostrata 1 0 Carditopsis smithi 2 14 4 7 2 Chione grus 1 1 2 2 10 I 5 2 8 3 Crassinella lunulata 13 9 42 I6 30 6 27 25 22 I6 I8 I9 Crenella divaricata 6 2 3 1 2 2 1 Cyclopecten nanus 3 7 9 2 3 5 1 5 3 Ervilia concentrica 1 I 1 0 Glans dominguensis 1 1 4 2 1 Gouldia cerina 5 I 5 11 7 9 23 4 8 3 12 12 8 Hiatella arctica 1 1 2 1 5 1 Limopsis sulcata I 0 Linga amiantus 2 1 0 Lucina nassula 5 2 3 1 1 Montacuta triquetra I 3 3 2 1 Musculus latera/is 1 1 2 1 2 1 Nucula proxima I 1 1 0

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Appendix 18: (Continued) Nuculana concentrica 2 9 3 15 2 Parastarte triquetra 1 0 Parvilucina blanda 4 0 Parvilucina multilineata 62 9 46 1 39 2 4 12 10 1 16 Pteromeris perplana 1 1 0 Tel/ina versicolor 4 1 3 4 3 1 Varicorbula op e rculata 7 10 29 2 1 16 3 6 Verticordia ornata 1 2 2 3 1 Vesicomya pilula 2 3 0 TOTAL BIVALVES 104 13 58 75 140 48 1 1 5 55 50 56 55 48 68 GASTROPODS 0 Acteocina candei 1 2 1 1 1 1 1 A lvania auberiana 1 2 16 1 10 34 7 17 11 15 9 10 Aorotrema pontogenes 1 1 1 0 Arene tricarinata 1 1 2 0 Bittium varium 1 0 Caecum bipartitum 1 7 1 3 2 1 Caecum clava 1 0 Caecum cubitatum 15 1 17 25 55 6 24 7 72 5 1 9 Caecum floridanum 3 1 2 5 1 Caecum imbricatum 2 3 4 3 1 Caecum nitidum 1 1 1 0 Caecum plicatum 3 1 0 Caecum pulchellum 10 40 66 4 22 117 47 54 18 51 104 4 4 Caecum torqu et um 1 7 1 1 Ca/yptraea centra/is 1 1 0 Cyc/ostremiscus cubanus 1 2 8 1 2 14 1 1 2 13 6 4 Cyclostremiscus jeannae 1 0 Finella dubia 17 17 35 37 1 6 13 17 4 12 Granulina ovuliformis 2 0 Mar_ginella /avalleeana 1 0 Margine/la sp. 1 2 1 1 0

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Appendix 18: (Continued) Natica pusi/la 5 4 3 I 2 3 2 2 Odostomia didyma 6 2 I 4 I Parviturboides interruptus I 0 Pyrunculus caelatus 2 I I 0 Retusa sulcata 3 I 3 I 2 I I Rissoina sp. 6 3 I Rissoina striatocostata 2 0 Seila adamsi I 0 Teinostoma biscaynense I 0 Teinostoma incertum 2 I 4 3 I I I Trico/ia thalassico/a I 2 I I 3 I Volvulella persimi/is 3 2 5 2 I 2 4 1 2 TOTAL GASTROPODS 62 51 119 61 93 202 157 87 70 26 183 152 105 SCAPHOPODS Cadulus iota 2 3 0 TOTAL SPECIMENS/SAMPLE 166 64 177 138 233 250 272 142 123 82 238 200 174

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....... .;:. 00 Appendix 19: Micromolluscan Production and Species List 1974 Transect I STATION NO. 13 14 15 16 17 18 19 20 DEPTH (METERS) 44 38 38 43 31 33 34 30 NO. SPECIMENS x 104 /m2 28.9 17.1 47.4 9.1 25.9 45.7 9.2 29.9 NO. LIVING SPECIMENS x 102 /m2 17.3 10.3 28.4 5.4 15.5 27.4 5.5 17.9 CALCIMASS in g CaC03 m-2 6.28 3.72 10.31 1.97 5.63 9.94 2.00 6.49 CaC03 PRODUCTION RATE in g CaC03 m-2 yr-1 62.78 37.23 103.09 19.69 56.31 99.40 20.00 64.93 BIVALVES Abra aequalis 4 Abra lioica 5 2 Astarte nana 1 1 Cardiomya ornatissima 1 1 Cardiomya perrostrata 1 1 Carditopasis smithi 1 2 Chionegrus 5 2 2 3 Crassinella lunulata 49 3 8 11 5 6 9 1 Crenella divaricata 4 1 2 3 2 CJ!clopecten nanus 4 2 Dacrydium vitreum 1 1 Glans dominf(Uensis 1 Gouldia cerina 21 4 13 7 6 20 9 19 Hiatella arctica 3 1 Linga amiantus 1 Musculus latera/is 2 1 1 2 1 Nucula proxima 2 1 4 21 22 STA. 30 42 36.3 27.3 46.4 28.7 16.4 27.8 17.2 5.94 10.09 6.24 59.39 100.93 62.38 0.4 0.7 0.2 0.2 0.2 0.3 1 1.3 5 9.7 1 3 1.6 1 0.7 0.2 0.1 16 18 13.3 1 0.5 1 0.2 3 1 1 3 1.1

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Appendix 19: (Continued) Parvilucina blanda 1 1 1 1 8 2 1 1.5 Parvilucina multilineata 18 56 128 2 85 133 1 90 64 118 69.5 Tel/ina versicolor 1 3 2 1 1 3 1.1 Thyasira trisinuata 1 0.1 Varicorbula opercula/a 7 5 33 3 9 31 16 17 19 14 TOTAL BIV AL YES 119 74 194 28 118 200 22 145 103 176 117.9 GASTROPODS Acteocina candei 1 3 1 3 1 0.9 Alvania auberiana 15 1 4 2 1 2.3 Arene tricarinata 3 1 1 2 1 0.8 Caecum bipartitum 12 27 30 60 2 48 58 92 32.9 Caecum clava 1 0.1 Caecum cubitatum 14 16 74 6 20 37 2 4 14 36 22.3 Caecum imbricatum 2 4 2 2 1 Caecum pulchellum 44 7 2 10 21 5 2 9.1 C)lclostremiscus cubanus 4 2 1 1 1 2 1.1 Finella dubia 4 8 26 8 7 16 5 4 8 13 9.9 Nannodiella melanitica 1 1 1 0.3 Natica pus ilia 3 2 2 2 2 4 1.5 Odostomia didyma 1 0.1 Parviturboides interruptus 1 0.1 Retusa sulcata 2 2 5 1 1 1 1 1.3 Seila adamsi 1 0.1 Teinostoma incertum 1 0.1 Volvulella persimilis 1 2 1 2 1 2 0.9 TOTAL GASTROPODS 85 47 141 36 65 123 43 66 90 152 84.8 TOTAL SPECIMENS/SAMPLE 204 121 335 64 183 323 65 211 193 328 202.7

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Appendix 20: Micromollusc Species List 1975-1976 Bivalves Abra lioica Amygdalum papyrium Barbatia dominguensis Bathyarca glomerula Botula fasca Cardiomya perrostrata Cardiomya sp. Corbula cymella Crassinella lunata Crassinella lunulata Crenella divaricata Cyc/opecten nanus Cyc/opecten nanus Cymatoica orienta/is Diplodonta punctata Erycina emmonsi Eucrassatel/a speciosa Gastrochaena hians Gouldia cerina Laevicardium pictum Limopsis cristata Limopsis sulcata Linga sombrerensis Lucina radians Lyonsia hyalina jloridana Nemocardium peramabile Nucinella adamsi Nuculana acuta Nuculana carpenteri Parvilucina multilineata Pitar simpsoni Plicatula gibbosa Semele nuculoides Solemya velum Tel/ina aequistriata Tel/ina versicolor Thyasira trisunuata Varicorbula operculata Verticordia ornata Gastropods Acteocina candei Atys riiseana Brachycythara barbarae Caecum bipartitum Caecum cubitatum Caecum pulchel/um Calyptraea centra/is Cerithiopsis crystal/inurn Cerithium atratum Crepidula fornicata Eulima bifasciatus Eulimostrica hemphilli Finella dubia 0/ive//a sp. Philine sagra Polyplachophora Acanthochitona pygmaea Ischnochiton papillosus Aplacophora Chaetoderma sp. Scaphopods Cadulus parvus Cadulus quadridentatus Dentalium bartletti Dentalium sp. Dentalium texasianum 150

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Appendix 21: Micromolluscan Production 1975-1976 Station Depth Sed. Summer Percent Fall Winter Annual Average Calcimass CaC03 PRODUCTION No. Type Density Living Density Density Living RATE m spec/m2 spec/m2 spedm2 spec .1m2 g CaC03m 2 g CaC03 m2 yr 23 11 f 1052.19 2.17% 631.31 631.31 771.60 2.80 27.97 24 18 f 210.44 0.68% 420.88 210.44 280.58 1.02 10.17 25 37 m 1052.19 0 .38% 841.75 1262 63 1052.19 3 .81 38.14 26 53 c 0.00 0.00% 210.44 420 88 210.44 0.76 7.63 27 90 c 0.00 0.00% 420.88 210.44 210.44 0.76 7 63 28 168 vf 210.44 0 .65% 631.31 0 .00 280 .58 1.02 I 0.17 STATION AVERAGE 63 420.88 0.65% 526.09 455.95 467.64 1.70 16.95 29 19 f 210.44 0.13% 420.88 420.88 350 .7 3 1.27 12.71 30 31 s 210.44 0.09% 1052.19 420 88 561.17 2.03 20.34 ...... 31 34 1683.50 0.69% 210.44 631.31 841.75 3 05 30.51 Vl s ...... 32 37 vf 0.00 0 .00% 1473.06 6523.57 2665.54 9.66 96.63 33 43 c 420.88 0.16% 0.00 210.44 210 .44 0 76 7 63 34 189 vf 0.00 0.00% 0.00 0.00 0 .00 0 00 0.00 STATION AVERAGE 59 420.88 0.18% 526.09 1367.85 771.60 2.80 27 .97 35 176 s 0.00 0.00% 0.00 420.88 140.29 0 .51 5.09 36 43 sf 0.00 1473.06 736.53 2.67 26.70 37 38 sf 841.75 210.44 526 .09 1.91 19.07 38 35 sf 1262.63 0 85% 1473 .06 210.44 982 .04 3.56 35.60 39 29 vf 1893.94 2 86% 1052.19 7786.20 3577.44 12.97 129.68 40 20 m 0.00 0.00% 1262.63 420.88 561.17 2.03 20.34 STATION AVERAGE 57 789.14 0 93% 771.60 1753 65 1087.26 3.94 39.41 1975-1976 AVERAGE 60 543.63 0.58% 607.93 1192.48 775.50 2.81 28.11

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Appendix 22: Archived Macromolluscs 1974 BIVALVES Aequipecten muscosus Americardia media A nadara jloridana Area imbricata Area zebra Barbatia domingensis Callista eucymata Chama macerophylla Chlamys benedicti Glycymeris americana Hiatella arctica Laevicardium sp. Lithophaga aristata Mercenaria campechiensis Pecten ravenel; Spondylus ictericus Tel/ina listeri Tel/ina simi/is Trachycardium sp. GASTROPODS Anachis sp. Aplysidae (Family) Calliostomas jujubinum Calliostomas pulchrum Cerithium litteratum Conus spurius at/anticus Coralliophila caribaea Crucibulum auricula Cymatium rubeculum occidentale Diodora sayi Engina sp. Fascia/aria /ilium Favartia cellulosa Marginella sp Modulus modulus Murex jlorifer dilectus Murex recurvirostris rubidus Pleuroploca gigantea Polinices sp. Simnia uniplicata Strombus gigas Triphora decorata Turbo castanea Turritella acropora Turritella exoleta Vermicularia lcnorri Vermicularia sp. Xenophora conchyliophora Transect I II X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X 152 POLYPLACOPHORA Acanthochitona sp. Ischnochiton sp. SCAPHOPODS Dentalium antalis taphrium Dorididae (Family) Rossia tenera CEPHALOPODS Octopus joubini Octopus sp. Transect I II X X X X X X X

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Appendix 23: Archived Dominant Macromolluscs 1975-1976 Transect Transect I IIIli I IIIli Bivalves Gastropods Abra /ioica X X X Acteocina candei X X Amygdalum papyrium X X Atys riiseana X X Barbatia dominguensis X X Brachycythara barbarae X Bathyarca glomerula X Caecum bipartitum X X Botula fasca X Caecum pulchellum X X Calyptraea centra/is X Cerithiopsis crystal/inurn X X Cardiomya perrostrata X Cerithium atratum X Cardiomya sp. X Crepidula fornicata X Corbula cymella X X Eulima bifasciatus X Crassinella lunulata X X X Eulimostrica hemphilli X Crenella divaricata X Finella dubia X X Cyclopecten nanus X X 0/ivella sp. X Cymatoica orienta/is X Philine sagra X Diplodonta punctata X X Erycina emmonsi X Scaphopods Eucrassatella speciosa X Cadulus cubitatum X Gastrochaena hians X X Cadulus parvus X Gouldia cerina X Cadulus quadridentatus X Laevicardium pictum X Dentalium bartletti X Limopsis cristata X Dentalium sp. X X Limopsis sulcata X Dentalium texasianum X Linga sombrerensis X Tel/ina aequistriata X Lucina radians X Lyonsia hyalina jloridana X X Nemocardium peramabile X Nucinella adamsi X Nuculana acuta X X Nuculana carpenteri X Parvilucina multilineata X X X Pitar simpsoni X X Plicatula gibbosa X Semele nuculoides X X Solemya velum X X Tel/ina versicolor X X X Thyasira trisinuata X X X Varicorbula operculata X X X Verticordia ornata X Polyplacophora Acanthochitona pygmaea X Chaetoderma sp. X X Ishnochiton papillosus X X 153

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Appendix 24: Macromolluscan Production 1974 Station Depth Biomass Calcimass Production No. Rate in meters gl 2 m m in g CaCOim2 in g CaC03/m2/yr 1 37 7.32 4.54 4.54 3 44 0.65 0.40 0.40 4 53 1.93 1.19 1.19 5 37 41.89 25.97 25.97 6 36 20.30 12.59 12.59 7 40 0 29 0.18 0.18 9 48 2.00 1.24 1.24 10 54 14.85 9.21 9.21 11 37 1.02 0.63 0.63 12 34 4.66 2.89 2.89 Transect II Average 42 9.49 5.88 5.88 13 44 6.36 3.94 3.94 14 38 4.63 2 87 2.87 15 38 0.16 0.10 0.10 17 31 0.98 0.61 0.61 18 33 2.60 1.61 1.61 19 34 1.79 1.11 1.11 20 30 0.36 0.22 0.22 21 30 0.52 0 32 0.32 22 42 0.28 0 .18 0.18 Transect I Average 36 1.96 1.22 1.22 Avg. 39 5.93 3.67 3.67 154

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Appendix 25: Macromolluscan Production 1975-1976 Station Depth Biomass Calcimass Production Density No. Rate meters g! 2 2 m m in g CaCOim g CaCO/m2/yr Spec/m 2 23 11 175.11 108.57 108.57 293 24 18 0 96 0.60 0.60 143 25 37 3.13 1.94 1.94 185 26 53 3.24 2.01 2.01 111 27 90 0.96 0.60 0.60 70 28 168 0.4 1 0.25 0.25 76 Transect I Average 63 30 64 18 99 18 99 146 29 19 6.29 3.90 3.90 311 30 31 3.29 2.04 2.04 313 31 34 1.69 1.05 1.05 348 32 37 8 02 4.97 4.97 880 33 43 2.02 1.25 1.25 148 34 189 0.74 0.46 0.46 72 Transect II Average 59 3 68 2 28 2.28 382 Average 61 17.16 10.64 10.64 2 64 155

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Appendix 26: Macromolluscan Production MMS 1992-1994 Station Depth Biomass Calcirnass Production Rate No. -2 g CaC03 m 2 g CaC03 m"2yr"1 mm mgm 41 IC 13 8.09 5 02 5 02 IC 13 1.13 0.70 0.7 0 IC 13 0.29 0.18 0.18 IC 13 6.71 4.16 4.16 Avg. 13 4.06 2 .51 2.51 42 ID 13 8.15 5.05 5.05 ID 13 20.48 12.70 12 70 ID 13 2.72 1.69 1.69 ID 13 0.97 0.60 0.60 ID 13 1.55 0 96 0 .9 6 ID 13 2 57 1.59 1.59 ID 13 27.89 17. 29 17.29 ID 13 2.72 1.68 1.68 Avg. 13 8.38 5 20 5.20 Avg. Site I 13 6.94 4.30 4.30 43 IIC 6 54 42 33.74 33 74 IIC 6 18.18 11.27 11.27 IIC 6 1.30 0.81 0 .81 Avg 6 24.63 15.27 15.27 44 IID 6 19 65 12. 19 12.19 liD 6 27.48 17. 04 17.04 liD 6 216.49 134.22 134.22 liD 6 122.38 75.87 75.87 liD 6 61.39 38.06 38.06 liD 6 2 84 1.76 1.76 Avg. 6 75.04 46. 52 46. 52 Avg. Site II 6 58 24 36.1 1 36 .11 45 IIIC 6 629.87 390 52 390 .5 2 46 IIID 6 2.37 1.47 1.47 HID 6 8 72 5.41 5.41 Avg 6 5 54 3.44 3.44 Avg. Site III 6 213.65 132.46 132.46 47 NC 6 3.80 2.36 2.36 48 ND 6 0.63 0.39 0.39 ND 6 1.07 0.66 0 66 Avg 6 0.85 0.53 0.53 Avg. SiteN 6 1.83 1.14 1.14 MMSAvg. 7.25 94 02 58.29 58.29 156

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Appendix 27: Echinoderm Species List Transect I Transect II 1975-1976 MMS Asteroidea Anthenoides piercei X Astropecten articulatus X Astropecten cingulatus X Astropecten comptus X X X Astropecten duplicatus X X X Astropecten nitidus X X Astropectinidae X Astroporpa annulata X Coscinasterias tenuispinus X Echinaster modestus X Echinaster sp. X X Goniaster tesselatus X X Luidia alternata X X Luidia clathrata X X X Luidia elegans X Narcissia trigonias X Oreaster reticulatus X Crinoidea Comactina meridiana/is X Echinoidea Araeosoma violaceum X Arbacia punctulata X X Brissopsis elongata X Clypeaster (durandi?) X X Clypeaster ravenelli X Clypeaster subdepressus X Diadema antillarum X Diadema sp. X Encope michelini X X Eucidaris tribuloides X X Lytechinus variegatus X X X X Mel/ita X Mel/ita tenuis X Meoma ventricosus X Moira X Moira atropos X Plagiobrissus X 157

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Appendix 27: (Continued) Transect I Transect II 1975-1976 MMS Holothuroidea Holothuria princeps X Holothuria sp. X Holothuroidea A X Holothuroidea B X Holothuroidea C X Holothuroidea D X Holothuroidea E X Leptosynapta X Leptosynapta crassipatina X Pseudothyone belli X Stichopus sp. X X Thyonella gemmata X Ophiuroidea Amphiodia X Amphiodia planispina X Amphiopholis X Amphipholis squamata X ? Amphiura fibulata X Astrophyton muricatum X Axiognathus squamata X Hemipholis elongata X Micropholis atra X Ophiacantha A (bidenta) X Ophiocoma sp. X X Ophioderma appressum X Ophioderma brevispinum A X Ophioderma brevispinum B X X Ophioderma cinereum X Ophioderma sp. X Ophiolepis elegans X X X X Ophionereis olivacea X Ophionereis reticulata X Ophiophragmus X Ophiophragmus wurdemani X Ophiostigma isacanthum X Ophiothrix angulata X X X Ophiothrix lineata X Ophiothrix suensonii A X Ophiothrix suensonii B X Ophiozona impressa X 158

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Appendix 28: Echinoderm Production 1974 and 1975-1976 Biomass Carbonate Station Depth Wet Weight Calcirnass Production inm in g m-2 gCaC03 m-2 g CaC03 m-2 yr-1 1 37 5.56 1.33 0.35 3 44 34.79 8.35 2.17 4 53 2.25 0.54 0.14 5 37 0.74 0.18 0.05 6 36 13.47 3.23 0.84 7 40 0.56 0.13 0.03 9 48 0.30 0.07 0.02 10 54 0.60 0.14 0.04 11 37 5.37 1.29 0.34 12 34 0.19 0.04 O.Ql Transect II A vg 42 6 38 1.53 0.40 13 44 0.18 0.04 0.01 14 38 230.45 55.31 14 .38 15 38 1.12 0.27 0.07 17 31 2.29 0.55 0.14 18 33 0.10 0.02 O.Ql 19 34 0.02 O.Ql 0.00 20 30 0.00 0.00 0.00 21 30 0.04 O.Ql 0.00 22 42 0.04 O.ol 0.00 Transect I A vg 36 26.03 6 25 1.62 1974 Avg 39 15.69 3.77 0.98 24 18 145.04 34.81 9.05 25 37 3.99 0.96 0.25 26 53 5.23 1.26 0.33 27 90 0.46 0.11 0.03 28 168 0.07 0.02 0.00 Transect I A vg. 73 30.96 7.43 1.93 29 19 51.16 12.28 3.19 30 31 0.07 0 02 0.00 31 34 0.07 0.02 0.00 32 37 4.88 1.17 0.30 33 43 3.60 0.86 0.22 34 189 0.29 0 07 0.02 Transect II A vg. 59 10.01 2.40 0.62 1975-1976 65 19.53 4.69 1.22 Avg. Shelf 49 17.10 4 10 1.07 159

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Appendix 29: Echinoderm Production MMS 1992-1994 Station Depth Site Biomass Calcimass Carbonate No. No. Wet Weight Production inm mgm in g CaC03 m -2 in g CaC03 m -2 yr -1 41 13 I 11.03 2.65 0.69 42 13 I 19 76 4 74 1.23 43 6 II 38 05 9.13 2. 3 7 44 6 II 4 .60 1.10 0 2 9 45 6 III 1.13 0.27 0 0 7 46 6 III 1.62 0 .39 0. 10 47 6 IV 207 85 49. 88 1 2 9 7 48 6 IV 2.91 0 70 0.18 MMSA vg. 35 87 8 .61 2.24 160

PAGE 171

Appendix 30: Dominant Archived Calcareous Algae I II FMG I II m Chlorophyta Avrainvillea asarifolia X X Halimeda discoidea X X Halimeda favulosa X X Halimeda opuntia X X Halimeda sp. Halimeda tuna X X Udotea conglutinata X Udotea cyathiformis X Udotea flabellum X X Phaeophyta Padina profunda X Padina vickersiae X Rhodophyta Amphiroa fragillisima X Fosliella atlantica X X Fosliella farinosa X Galaxaura obtusata X X Galaxaura sp. X Galaxaura squalida X Jania adherens X Jania capillacea X X Lithothamnion incertum X X Lithothamnion occidentale X X Lithothamnion sejunctum X Lithothamnion syntrophicum X Cyanophyta Anacystis acruginosa X Calothrix crustacea X Entophysalis conferta X Microcoleus lyngbyaceus X Microcoleus vaginatus X Schizothrix calcicola X Spirulina subsalsa X 161

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Appendix 31: Archived Hard and Soft Corals 1974 Transects 1975-6 Transects I II II III Octocorallia Bebryce grandis X X Bebryce parastellata X X X Diodogorgia nodulifera X X Ellisel/a barbadensis X Eunicea ca!ycu!ata X X Eunicea knighti X Leptogorgia eurya/e X Leptogorgia medusa X Lophogorgia cardinalis X Lophogorgia hebes X Muricea e/ongata X X Muricea laxa X X Neospongodes agassizii X Paramuric ea sp. X Plexaura flexuosa X Plexau r e lla fusifera X Pseudop/exaura porosa X R e nilla mulleri X Villogorgia nigrescens X Scleractinia Agaricia agaricites X Cladocora arbuscula X X X X Cladocora debilis X X Dichacaenia stellaris X X Dichacaenia stokesii X Isophyllia sinuosa X Madracis asperula X Madracis decactis X X X Manicina areolata X Millepora alciconzis X Oculina diffusa X X X Ocu/ina tenella X X Parac\athus dejilippi X X X Phyllangia americana X X Porites branneri X Porites divaricata X X Scolym ia cubensis X Sco/ymia la cera X Siderastrea radians X So/ e nastrea h yades X X Steplzanocoenia miclze lini X X X 162

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Appendix 32: Archived Serpulid Polychaetes Family: Serpulidae (Calcareous Tube Worms) Apomatus similes Apomatus sp. Crucigera websteri Dexiospira spieillum Ditrupa arietina Ficopomatus n macrodon Filograna huxleyi Filograna implexa Filograna sp. Hydroides bandaensis Hydroides bispinosus Hydroides caribensis Hydroides crucigera Hydroides dianthus Hydroides elegans Hydro ides floridanus Hydroides gairacensis Hydroides heteroceros Hydroides lunulifera Hydroides microtis Hydroides norvegica Hydroides parvus Hydroides protulicola Hydroides sceptrifer Hydroides sp. nov. Hydroides spongicola Hydroides uncinata Josephella marenzellerei Metavermilia multecristata Metavermilia sp. A 163 Neovermilia capensis Placostegus incomptus Placostegus tridentatus Pomatoceros americanus Pomatoleios caerulescens Pomatoleios sp. nov. Pomatostegus Protula balbaoensis Protula diomedae Protula sp. Protula tubularia Pseudovermilia fuscostriata Pseudovermilia multispinosa Pseudovermilia occidentalis Pseudovermilia sp. Rhodopsis pusilla Salmacine dysteri Sclerostyla ctenactis Serpula lobiancoi Serpula c.f. massiliensis Serpula sp. Serpula vermicularis Spirobranchus giganteus Spirorbis corrogatum Vermiliopsis annulata Vermiliopsis biformis Vermiliopsis infundibulum Vermiliopsis multiannulatum

PAGE 174

Appendix 33: Florida Middle Ground Production C/.) ....... = Q) E :.a ell ell ell Q) (.) (.) ....... (/.) ell C/.) C/.) ell E 1-o ::l ::l = .5 a 0 0 ell Q) ell E E (.) "0 :.a = ...c 0 C/.) 0 Q) 9 0 ::l = Q) 0.. ...... ell 1-o 1-o ell Cil Cil ....... 0.. u (.) (.) :.a : ell Q) 1-o ell 0 ..::fl 1-o ...... ....... Cl 0 : ::E ::E (.) 0 0 (/.) --< u u f-c Florida Middle Ground Stations 3 44 93 18.90 54.47 0.40 54.87 2.17 25.00 3 00 0 50 104.44 5 37 64 26.20 71.70 2 5 97 97.67 0 .05 25.00 3 00 0 .50 1 52.42 6 36 98 1 2 .10 76.93 12. 59 89.52 0 84 25. 00 3.00 0 50 130 96 7 40 84 5 2 .90 83. 70 0 .18 83.88 0 .03 25. 00 3 00 0 50 165.31 8 42 93 24.30 43. 70 8.41 52.11 0.62 25. 00 3.00 0.50 105.53 12 34 77 11.50 61.54 2.89 64.43 0 .01 25. 00 3.00 0.50 104 44 36 43 10 22.02 26.70 8.41 35.11 0.62 25. 00 3.00 0.50 86.25 37 38 96 35. 70 19. 07 8.4 1 27.48 0 62 25. 00 3.00 0.50 92 30 38 35 80 8.57 35.60 8.41 44.0 1 0.62 25. 00 3 00 0.50 81. 70 FMG I Avg. 39 77 23.58 52. 60 8.41 61.01 0 62 25. 00 3.00 0.50 113.71 Stations Surrounding Florida Middle Ground 1 37 78 51.20 51.08 4.54 55. 62 0 .35 2 .50 0 30 0 00 1 09 97 2 45 86 46.50 19.69 3 32 23.01 0 .83 2 .50 0 30 0.00 73.14 4 53 78 37.40 42.47 1.19 43.66 0.14 2.50 0 30 0.00 84.00 9 48 90 58. 1 0 37.85 1.24 39 09 0.02 2 .50 0 30 0.00 100.01 10 54 92 86.14 25.23 9.2 1 34.44 0 04 2 .50 0 30 0.00 123.42 11 37 91 31.60 73. 24 0.63 73.87 0 34 2 50 0.30 0.00 108.61 46 86 51.82 41. 59 3 36 44 .95 0 28 2.50 0.30 0.00 99.86 Average of Stations without Florida Middle Ground 42 7 1 41.20 36 .93 17. 56 54.49 1.42 3 .52 0.42 0.00 101.04 Central West Florida Shelf Station Average 42 72 37 .89 39.87 15. 84 55.71 1.27 7.55 0.90 0 09 103.41 164

PAGE 175

Appendix 34: Total Carbonate Production on the Central West Florida Shelf ] "' "' s u u u tl) "' "' "' e .s 1 ..= ..= .s 0 ] "' u = 8 e u '"0 '"0 .;3 8 "' 0 u 0 0 ..= s u ] : .:: 0.. "' .... u 0 "' 3 s .... "' ..c: u 0 u 0 0 tl) 0 :::R j;.I., l:.t.l < u u E-< 0 1 37 78 51.20 51.08 4.54 55.62 0.35 2.50 0.30 0.00 109.97 2 45 86 46.50 19.69 3.32 23.01 0 .83 2.50 0.30 0.00 73.14 3 44 93 18 90 54.47 0.40 54.87 2.17 25.00 3.00 0.50 104.44 4 53 78 37.40 42.47 1.19 43.66 0.14 2.50 0 30 0.00 84.00 5 37 64 26.20 71.70 25.97 97.67 0.05 25.00 3.00 0.50 152.42 6 36 98 12.10 76.93 12.59 89.52 0.84 25.00 3 00 0.50 130.96 7 4 0 84 52.90 83.70 0.18 83.88 0.03 25. 00 3.00 0.50 165.31 8 42 93 24 30 43.70 8.41 52.11 0 62 25 00 3 00 0.50 105.53 9 48 90 58.10 37.85 1.24 39.09 0.02 2.50 0.30 0 00 100 .01 10 54 92 86.14 25.23 9.21 34.44 0.04 2.50 0.30 0.00 123.42 11 37 91 31.60 73.24 0.63 73.87 0.34 2.50 0.30 0.00 108.61 12 34 77 11.50 61.54 2.89 64.43 0.01 25 00 3.00 0.50 104.44 42 85 38.07 53.47 5.88 59.35 0.45 13.75 1.65 0.25 113.52 13 44 92 87.61 62 78 3.94 66.72 0.01 2.50 0.30 0 00 157.14 14 38 90 61.44 37.23 2.87 40.10 14.38 2.50 0 30 0.00 118.72 15 37 89 39.14 103.09 0.10 103.19 0.07 2.50 0.30 0 00 145.20 16 43 97 5.99 19.69 3.32 23 .01 0.83 2.50 0.30 0.00 32.63 17 31 63 123.63 56.31 0.61 56.92 0.14 2.50 0 30 0.00 183.49 18 33 88 52.46 99.40 1.61 101.01 0.01 2.50 0.30 0.00 156.28 19 34 96 9.93 20 00 1.11 21.11 0.00 2.50 0.30 0.00 33.84 20 30 84 65. 34 64.93 0.22 65.15 0.00 2.50 0.30 0.00 133.29 21 30 84 120.04 59.39 0.32 59.71 0.00 2.50 0.30 0.00 182.55 22 42 81 92 03 100.93 0.18 101.11 0.00 2.50 0.30 0.00 195.94 36 86 65 76 62.38 1.43 63.80 1.54 2.50 0.30 0.00 133.91 165

PAGE 176

Appendix 34: (Continued) .l!l s:: s ;a "' "' .s u u til "' "' "' s s .2 .2 E 5 a 0 0 "' "' .e s s u "0 ;a s:: a "' 0 0 ;S 0 0 .2 dJ "' .... dJ 0.. c.. u "' .... u "' c;j E c;j .s .... 0 bO .... dJ 0 "' u 0 0 til 0 "

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