Ecological Functions of 

Nearshore Hardbottom Habitats 

in East Florida: A Literature Synthesis 

June 2009 

Prepared for: 

Bureau of Beaches and Coastal Systems 

Florida Department of Environmental Protection 

3900 Commonwealth Boulevard, M.S. 300 

Tallahassee, Florida 32399 

Prepared by: 

CSA International, Inc. 

8502 SW Kansas Avenue 

Stuart, Florida 34997 
Ecological Functions of 

Nearshore Hardbottom Habitats 

in East Florida: A Literature Synthesis 

June 2009 

Prepared for: 

Bureau of Beaches and Coastal Systems 

Florida Department of Environmental Protection 

3900 Commonwealth Boulevard, M.S. 300 

Tallahassee, Florida 32399 

Prepared by: 

CSA International, Inc. 

8502 SW Kansas Avenue 

Stuart, Florida 34997 

Authors: 

Kenyon C. Lindeman 

Department of Marine and Environmental Systems 

Florida Institute of Technology 

Melbourne, Florida 

Daniel A. McCarthy 

Department of Biology and Marine Science 

Jacksonville University 

Jacksonville, Florida 

Karen G. Holloway-Adkins 

East Coast Biologists, Inc. 

Indialantic, Florida 

David B. Snyder 

CSA International, Inc. 

Stuart, Florida 
Acknowledgments 

Acknowledgments

The genesis, funding, and review of this project were made possible by the Florida Legislature through 

the Florida Department of Environmental Protection Bureau of Beaches and Coastal Systems Beach 

Management Funding Assistance Program. The report was prepared under contract to PSBJ. We thank 

the project’s Technical Advisory Committee (TAC) for their assistance and editorial comments throughout 

the project. TAC members included Martin Seeling, Roxane Dow, and Vladimir Kosmynin (Bureau of 

Beaches and Coastal Systems), Cheryl Miller (Coastal Eco-Group Inc.), Don Deis (PBSJ), Jeff Beal and 

Dr. Robin Trindell (Florida Fish and Wildlife Conservation Commission), Terri Jordan (U.S. Army Corps of 

Engineers), and Jocelyn Karazsia (National Marine Fisheries Service). At CSA International, Inc. (CSA), 

we thank Melody Powell for proofreading and editing the entire document and Karen Stokesbury for 

coordinating document production. Keith VanGraafeiland and Chris Dean at CSA also assisted with 

aerial imagery and nearshore hardbottom acreage estimates. Staff from agencies and consulting 

companies, including Ken Banks (Broward County Department of Environmental Protection), Brian Flynn 

(Miami-Dade Department of Environmental Resources Management), Brian Walker (National Coral Reef 

Institute, Nova Southeastern University Oceanographic Center), Janet Phipps and Carmen Vare (Palm 

Beach County Department of Environmental Resources Management), and Quin Robertson (Coastal 

Planning and Engineering, Inc.), also provided estimates of hardbottom acreage. We gratefully 

acknowledge Julia Goodman (Jacksonville University) for the hours she spent compiling lists of 

invertebrates encountered in the various studies we reviewed. We thank Gerry Pinto (Jacksonville 

University) for developing the draft of the arthropod section within the invertebrate chapter. Jon 

Norenburg and Mary Rice of the Smithsonian Institute provided significant input on nemertines and 

sipunculans, respectively. We thank Bailey Estrada (East Coast Biologists, Inc.) for many hours helping 

develop the figures and tables in the algae and marine turtle chapters, Larry Wood (Palm Beach Zoo) for 

sharing his hawksbill turtle research findings, and Mark and Diane Littler (Smithsonian Institute) for 

permission to reference their functional form tables and concepts. We thank Scott Hurley (Harbor Branch 

Foundation) and Kirsten Jones (Gumbo Limbo Nature Center) for allowing use of their photographs to 

better illustrate some nearshore hardbottom habitats. We also thank Stacy Prekel and Jessica Craft of 

Coastal Planning and Engineering, Inc. for graciously provided data from their monitoring projects. 

Finally, we thank Lance Jordan (Nova Southeastern University Oceanographic Center) and Grant 

Gilmore (Estuarine, Coastal, and Ocean Science, Inc.) for reviews and comments on several chapters. 

Suggested Citation: 

CSA International, Inc. 2009. Ecological functions of nearshore hardbottom habitat in east Florida: A 

literature synthesis. Prepared for the Florida Department of Environmental Protection Bureau of 

Beaches and Coastal Systems, Tallahassee, FL. 186 pp + apps. 
Executive Summary ES-1 

Executive Summary

Given the complex policy questions involving hardbottom burial and mitigation, the consensus need for 

ecosystem-based management, and the large but unsynthesized literature base, this study responds to 

four issues: 1) What are specific ecological functions of nearshore hardbottom (NHB) (0 to 4 m depth)?, 

2) What is happening to the coastal ecosystem (broader than NHB) when NHB is lost due to 

nourishment?, 3) Is offshore hardbottom (OHB) (>6 m depth) able to compensate for the loss of NHB?, 

and 4) If the NHB habitat has specific properties and functions that cannot be compensated for by 

existing OHB habitat, what are the best mitigation alternatives for the loss of natural NHB? 

Very little depth-specific data exist for a majority of taxa throughout the region, which currently limits some 

of our ability to summarize core functional differences between NHB and OHB. However, a good deal of 

new information has been assembled and examined on shallow hardbottom structure and distributions 

(Section 1); the algal, invertebrate, fish, and turtle assemblages associated with shallow hardbottom 

(Section 2); and primary ecological functions of shallow hardbottom with a focus on latitudinal and depth 

variations, and implications for improved mitigation planning and implementation (Section 3). 

ALGAE, INVERTEBRATE, FISH, AND TURTLE ASSEMBLAGES 

Shallow hardbottom of southeast and east central Florida supports a sometimes diverse reef assemblage 

consisting of algae, invertebrates, fishes, and sea turtles. The fish and sea turtle components of the 

assemblage tend to be dominated by juvenile life stages. Patches of NHB exist in portions of Broward, 

Palm Beach, Martin, St. Lucie, Indian River, and Brevard Counties. Although these areas typically are 

separated by kilometers of sand, there are some areas where there is a semi-continuous hardbottom 

gradient from the beach into depths exceeding 6 m. Offshore, many mid-shelf areas are also dominated 

by expanses of sand, despite the variable occurrence of several mid-shelf reefs. Therefore, in many 

shallow areas, there are no natural habitats in the same or adjacent nearshore areas that can support 

equivalent abundances of early life stages of fishes and the invertebrates on which they largely feed. 

Absences of nursery structure can result in increased predation and lowered growth. In newly settled and 

juvenile stages, such conditions could foster demographic bottlenecks that ultimately result in lowered 

local population sizes (Jones, 1991). 

Algae 

The total number of recorded algal taxa, both identified and unidentified, is approximately 340, including 

277 known species. Potentially dominant genera are Ceramium, Dictyota, Hypnea, Laurencia, Gelidium, 

Caulerpa, Jania, and Ulva, depending on subregions and other factors. Sunlight converted into 

macroalgal tissue is directly eaten by as many as 20 genera of invertebrates, at least 14 genera of fishes, 

and the juvenile stage of the endangered green turtle, Chelonia mydas. 

Algae are a dominant driver of the food web and also contribute to shelter used by hundreds of species of 

invertebrates and fishes. All algal species contribute to oxygen and nutrient production. In terms of 

functional form groups, jointed-calcareous algae are most responsible for sediment enrichment and sand 

building. The abundance and diversity of macroalgae on NHB vary substantially with changes in latitude 

from Cape Canaveral to Miami Beach. 

Invertebrates 

Over 533 species of invertebrates have been identified along the east Florida coast, and there are likely 

to be more. The diversity of sessile species is greatest for cnidarians (~21%), bryozoans (~29%), and 

sponges (~19%). Some sessile groups may not be represented with such high diversity but can occur in 

very high biomass. A primary example is the sabellariid polychaete Phragmatopoma lapidosa, which can 
be very abundant along the mid- to north sections of the project area and is a habitat engineer that 

creates structure supporting high diversities of many other invertebrates. 

The most diverse and generally abundant motile invertebrates are arthropods and polychaetes, which 

represent 47% and 28%, respectively, of the total number of motile invertebrate species. Over 

100 species of crustaceans are extremely abundant, especially on worm reef-dominated hardbottoms. 

These include crabs, stomatopods, shrimp, isopods, and amphipods. There are over 87 reported 

polychaete species that are likely to be very abundant, although more research is needed to confirm this. 

There are also fairly high numbers of gastropods, flat worms, ribbon worms, and echinoderms on these 

habitats. 

Primary ecological functional roles of invertebrates along the east Florida coast include 

1) shelter-enhancing organisms that increase local diversity of fishes and invertebrates, and 2) predators 

or prey in local food webs. Generally, the highest community biomasses along the mainland east Florida 

coast occur in hardbottom areas with higher abundances of sessile invertebrate species (some may be 

considered foundational or keystone contributors to the community) that enhance local shelter. Along the 

Florida coast, important shelter-enhancing taxonomic groups are hard and soft corals, sponges, 

tunicates, molluscs, barnacles, and polychaetes (i.e., P. lapidosa). However, their importance in this 

function may vary dramatically with depth and latitude. In terms of contribution to local food webs, 

important taxonomic groups are sponges, polychaetes, echinoderms, crabs, and shrimp. 

The loss of NHB would result in a significant reduction of a high biomass of invertebrates (sponges, 

corals, crabs, shrimp, worms, gastropods, bivalves) that could significantly change the nearshore food 

web. Before a dredging event, the nearshore community can be fairly diverse, consisting of a number of 

opportunistic (common) and longer-lived (often uncommon) species. The diversity contributes to a 

nearshore food web that consists of suspension feeders, herbivores, omnivores, and various levels of 

carnivores. However, after a dredging event occurs that results in high mortality and habitat elimination, 

there is likely to be a less diverse community consisting of more opportunistic species and fewer rare 

species such as corals. Further, conditions where there are significant reductions in shelter enhancement 

in these areas (via fewer sessile species) could reduce the number of small carnivore trophic levels and 

have possible adverse effects on the higher trophic levels within the assemblage. 

Natural recovery of hardbottom habitats depends on both re-exposure of substrate as well as the 

recruitment capability of sessile shelter-enhancing invertebrates which in turn likely affect associated 

motile invertebrate species. Although it requires further investigation, it appears that very few older adult 

motile invertebrates would migrate into the area of impact. Consequently, habitat recovery requires 

ample larval supply and recruitment of the shelter-enhancing species to facilitate what could be a slow 

recovery of many associated motile invertebrates. The slow recovery of crevice-dwelling species 

(e.g., small crabs, shrimp, worms) may in turn affect prey availability for many motile invertebrate 

predators and some fishes. Further, degradation in water quality (most likely via fine resuspension and 

chronic turbidity) may affect fitness and/or survival of suspension feeders. 

It is unlikely that OHB is able to compensate for the loss of NHB, particularly in the northern areas of the 

east Florida coast. Mitigation efforts to restore hardbottom habitats in some areas likely depend on 

recruitment capability of shelter-enhancing species such as P. lapidosa. Local hydrodynamics and the 

type, rugosity, and relief of construction materials must be considered for artificial reefs to maximize 

recruitment and subsequent recovery of the habitat. For instance, in the case of P. lapidosa, abundance 

decreases significantly in water deeper than 4 m. Even within the 0 to 4 m depth range, the highest 

abundances occur more towards the intertidal zone. This may be related to better feeding and 

reproductive opportunities and greater availability of suspended sand particles for tube-building in these 

more turbulent shallow waters. However, the mechanism by which worms settle and survive in these 

habitats is via large-scale hydrodynamics that affect larval supply and various biotic/abiotic factors that 

affect juvenile mortality. It is likely (though it requires further investigation) that settling P. lapidosa larvae 

may touch bottom in deeper waters, but actively postpone metamorphosis and choose to settle when they 

encounter shallower waters (McCarthy, 2001). 

Executive Summary ES-2 
Consequently, if the goal is to mitigate for loss of worm reefs (and associated organisms), then structures 

deployed in the OHB would not be successful unless chemical cues, hydrodynamics, or other 

environmental factors could be created that accurately mimic the shallower waters and induce larvae to 

settle. Further, if settlement does occur at these depths, growing juveniles must survive, and recruitment 

of associated species must occur to properly mitigate for lost habitat. 

Fishes 

Currently, 257 species of fishes are recorded from NHB habitats. Species composition for the NHB 

ichthyofaunal assemblage is similar to that from shallow coral reef faunas from the region. Empirical 

information is highly limited on the amount of connectivity between shallow patches of hardbottom and 

deeper reefs for fishes and invertebrates of east Florida. The available information suggests that diverse 

fish species that ontogenetically migrate into deeper water can use NHB during early life stages. The 

physical environment is subject to wind and wave disturbance of sometimes high levels at intermediate 

temporal and spatial scales, and NHB assemblages are disturbance-mediated. If the NHB in use is close 

enough to intermediate hardbottom (IHB) or OHB, some of these species, depending on many changing 

ecological drivers (predation, growth, larval recruitment), could ontogenetically migrate into deeper 

habitats. Answers to the question “How close is close enough?” will vary by species, site, and time. 

Tagging studies using either conventional or acoustic methods that compare cross-shelf movements of 

fishes associated with isolated bands of NHB with movements of fishes associated with hardbottom 

extending continuously across the depth gradient from NHB to OHB would provide valuable insights. 

Some differences among ichthyofaunal assemblages are present between the southern and northern 

areas of mainland southeast and east central Florida in terms of the most abundant species. This 

information suggests there is a southern NHB ichthyofauna and a less diverse northern fauna, particularly 

north of the deflection of the Gulf Stream offshore. Potential explanations for such difference in dominant 

species within 100 to 200 km north-south distances may include temporal recruitment variations, NHB 

structural differences between the two areas, species reaching their biogeographic distributional limits 

between the northern and southern areas, and other factors. 

NHB structural differences probably exist between these areas as described above, but many of the more 

prominent species appear to be relatively opportunistic in their use of differing NHB and IHB microhabitat 

structure. At least three of the prominent species (black margate, sailors choice, and hairy blenny) are 

listed by Gilmore and Hastings (1983) as species whose Florida distributions are restricted to east 

Florida, not occurring in the Gulf of Mexico as congeners do. More research is required to parse the 

multiple factors and examine the above alternatives to better understand these and other north-to-south 

differences in fish assemblage pattern. 

Turtles 

Juvenile sea turtles of three species commonly associate with shallow hardbottom of mainland east 

Florida. Juvenile stages of the endangered green turtle, Chelonia mydas, can associate with shallow 

hardbottom for years, feeding on macroalgae and using structure for shelter. NHB certainly represents 

facultative habitat for juvenile green turtles that may utilize estuarine areas, as well. However, juvenile 

green turtle populations residing in bays and lagoons have shown high occurrences of disease and injury. 

As human populations and construction increase in Florida, nearshore reef habitat may play a 

comparatively more significant role in the recovery of these endangered species. There also are many 

interactions between turtles and an array of organisms that utilize shallow hardbottom. For example, 

hawksbill turtles Eretmochelys imbricata fragment sponges when foraging, creating space and habitat for 

other organisms, as well as increasing sponge growth through fragmentation. Globally, DNA evidence 

indicates juvenile marine turtles on NHB represent a mixed stock of progeny from the Caribbean, Mexico, 

Costa Rica, and Mediterranean nesting grounds. 

Executive Summary ES-3 
Executive Summary ES-4 

ECOLOGICAL FUNCTIONS AND MESOSCALE CONNECTIVITY 

Many species use NHB, IHB, and OHB as settlement and juvenile grow-out areas. The majority of the 

fish species are abundantly recorded from coral reefs, as are many other resident and transient 

organisms. For fishes, NHB assemblages are not only juvenile dominated, they are reef-species 

dominated. As the only natural habitat structure within a radius of 0.5 to 5 km of sand in any direction, 

NHB structural features potentially serve a variety of ecosystem functions, including settlement and 

nursery areas, spawning sites (for over 100 species of invertebrates and fishes), economic drivers, and 

recreational service providers. 

For sessile invertebrates, there appears to be a trend towards limited dispersal of larvae or juveniles 

along the east Florida coast. Of those species that reproduce sexually and asexually, many seem to 

propagate more via asexual reproduction. Recent molecular research supports this idea, as there 

appears to be restricted gene flow among Caribbean populations. Considerable information has been 

amassed regarding a variety of biotic and abiotic factors that contribute to retention in larvae. 

Developing better information on population connectivity can allow greater understanding of impact 

cascades at more refined spatial, temporal, and population scales. Efforts are underway to assign select 

species or families of primary groups to appropriate connectivity categories. Species with short larval 

periods show attributes of populations that could be categorized in patterns reflecting either 1) mesoscale 

dispersal structured by physics, resulting in several metapopulation substructures, or 2) diffusive dispersal 

with stepping-stone metapopulations. These connectivity patterns may change markedly above and 

below the St. Lucie deflection of the Florida Current. 

MITIGATION AND RESEARCH NEEDS 

Increasingly important and expensive questions involve the use of mitigation projects to offset impacts 

from large dredge and fill projects. Typically, these questions represent key steps in the finalization of 

permitting processes. Preliminary conclusions regarding the functional attributes of NHB and the 

application of that information to the evaluation of mitigation alternatives include the following: 

• Mitigation of NHB impacts by utilizing artificial reefs can often provide adequate replacement of lost 

structural habitat and therefore some structure- and trophic-associated ecological functions. 

• Not all mitigation reef designs and deployments will be successful in all situations, and site-specific 

factors must be considered. Different approaches and designs will be required for different locations 

in southeast and east central Florida. 

• For worm reef areas, ensuring successful local larval recruitment events is essential. To this end, 

project designs need to ensure that sediment size range is suitable to incoming recruits. 

• In terms of Issue 4, NHB functions from Brevard to mid-Palm Beach County will not be fully replaced 

with artificial reefs in deeper NHB, IHB, or OHB water depths because of the depth limits of the 

habitat engineering worm species P. lapidosa and the high numbers of associated species that 

depend on P. lapidosa structure for shelter. 

• A full conceptual ecological model (CEM) needs to be developed; this will be more efficient if a 

thorough food web draft is available. Much of the information to build a larger CEM is now available. 

• Artificial reef mitigation does not appear to remove or provide compensatory mitigation for the direct 

and indirect physiological and potential population scale impacts of turbidity and sediment 

resuspension. 

• The intermediate or long term water quality effects on shallow hardbottom from turbidity and 

sedimentation events, long-shore and cross-shelf, that result from dredge and fill projects (and 

relationships to natural turbidity levels) require examination. Without this type of information, 

responses to Issue 2 regarding whole ecosystem effects when NHB is buried or to Issue 3 regarding 

OHB compensation roles will be not be fully resolvable. 

• Approximately 340 algal taxa can associate with NHB, and most are subject to high mortality on a site 

that is filled. Mitigation reefs can provide new attachment sites for algae. Algal growth on mitigation 

reefs should correlate in part with water clarity. 
Table of Contents i 

Table of Contents

Page 

EXECUTIVE SUMMARY ........................................................................................................................ ES-1 

LIST OF TABLES ......................................................................................................................................... v 

LIST OF FIGURES ..................................................................................................................................... vii 

LIST OF ACRONYMS ................................................................................................................................. xi 

Section 1 Project Objectives and Nearshore Hardbottom within the 

East Florida Seascape 

CHAPTER 1 INTRODUCTION ..................................................................................................................1-1

1.1 INTRODUCTION AND OBJECTIVES ..........................................................................................1-1

1.2 ECOLOGICAL CONCEPTS AND TERMS ...................................................................................1-6

1.3 NEARSHORE, INTERMEDIATE, AND OFFSHORE HARDBOTTOM.........................................1-7

1.4 SPECIES: WINDOWS INTO ASSEMBLAGES? ..........................................................................1-8

1.5 EQUILIBRIUM AND SCALE .........................................................................................................1-9

1.6 ECOLOGICAL FUNCTIONS.........................................................................................................1-9

CHAPTER 2 SHALLOW HARDBOTTOM OF EAST FLORIDA AND THE REGIONAL SHELF 

SETTING...............................................................................................................................................2-1

2.1 INTRODUCTION...........................................................................................................................2-1

2.2 GEOLOGICAL AND BIOLOGICAL ATTRIBUTES OF NEARSHORE HARDBOTTOM...............2-2

2.3 DISTRIBUTION OF SHALLOW HARDBOTTOM IN EAST FLORIDA, MIAMI TO 

CANAVERAL ................................................................................................................................2-5

2.4 OFFSHORE HARDBOTTOM AREAS ..........................................................................................2-6

2.5 REGIONAL SETTING: THE FLORIDA CURRENT AND THE CONTINENTAL SHELF ............2-10

2.5.1 Currents and Upwelling Events......................................................................................2-10

2.5.2 Tides, Wind Events, Wave Activity, and Hurricanes......................................................2-11

Section 2 Organismal Assemblages of East Florida Shallow Hardbottom 

CHAPTER 3 MACROALGAE AND CYANOBACTERIA..........................................................................3-1

3.1 INTRODUCTION...........................................................................................................................3-1

3.1.1 Functional Form Groups ..................................................................................................3-1

3.1.2 Latitudinal Gradients ........................................................................................................3-3

3.1.3 Depth Gradients...............................................................................................................3-4

3.1.4 Algal Reproduction...........................................................................................................3-5

3.1.5 Functional Form Group Descriptions ...............................................................................3-7

3.2 SUMMARY..................................................................................................................................3-15

CHAPTER 4 INVERTEBRATES ...............................................................................................................4-1

4.1 INTRODUCTION...........................................................................................................................4-1

4.2 SESSILE SPECIES.......................................................................................................................4-1

4.2.1 Polychaetes......................................................................................................................4-1

4.2.2 Anthozoans ......................................................................................................................4-6

4.2.3 Sponges .........................................................................................................................4-11

4.2.4 Hydrozoans....................................................................................................................4-12
Table of Contents

(Continued)

Page 

Table of Contents ii 

4.2.5 Tunicates and Bryozoans ..............................................................................................4-13

4.2.6 Molluscs (Sessile) ..........................................................................................................4-14

4.2.7 Barnacles .......................................................................................................................4-17

4.3 MOTILE SPECIES ......................................................................................................................4-18

4.3.1 Crustaceans...................................................................................................................4-18

4.3.2 Gastropods and Other Molluscs ....................................................................................4-30

4.3.3 Echinoderms ..................................................................................................................4-33

4.4 SUMMARY..................................................................................................................................4-35

4.4.1 Diversity .........................................................................................................................4-35

4.4.2 Trophic Patterns and Functional Groups .......................................................................4-35

4.4.3 Latitudinal and Cross-shelf Distribution .........................................................................4-36

4.4.4 Reproduction and Life History........................................................................................4-36

4.4.5 Dispersal and Connectivity ............................................................................................4-37

4.4.6 Recruitment and Cross-shelf Habitat Use .....................................................................4-37

4.4.7 Economic and Recreational Value.................................................................................4-38

4.4.8 The Importance of Hardbottom to Invertebrates............................................................4-38

CHAPTER 5 FISH ASSEMBLAGES.........................................................................................................5-1

5.1 INTRODUCTION...........................................................................................................................5-1

5.2 DEMERSAL CARNIVORES .........................................................................................................5-3

5.2.1 Focal Families and Species .............................................................................................5-3

5.2.2 Shelter Use ......................................................................................................................5-4

5.2.3 Trophic Patterns...............................................................................................................5-8

5.2.4 Developmental Patterns and Cross-shelf Habitat Use ....................................................5-8

5.2.5 Latitudinal Distribution......................................................................................................5-9

5.3 CRYPTIC HARDBOTTOM RESIDENTS......................................................................................5-9

5.3.1 Focal Families and Species .............................................................................................5-9

5.3.2 Shelter Use ......................................................................................................................5-9

5.3.3 Trophic Patterns.............................................................................................................5-11

5.3.4 Latitudinal Distribution....................................................................................................5-11

5.4 COASTAL PELAGICS ................................................................................................................5-11

5.4.1 Focal Families and Species ...........................................................................................5-11

5.4.2 Shelter Use ....................................................................................................................5-12

5.4.3 Trophic Patterns.............................................................................................................5-14

5.4.4 Latitudinal Distribution....................................................................................................5-14

5.5 HERBIVORES AND OMNIVORES.............................................................................................5-15

5.5.1 Focal Families and Species ...........................................................................................5-15

5.5.2 Shelter Use ....................................................................................................................5-15

5.5.3 Trophic Patterns.............................................................................................................5-16

5.5.4 Developmental Patterns and Cross-shelf Habitat Use ..................................................5-17

5.5.5 Latitudinal Distribution....................................................................................................5-17

5.6 SUMMARY..................................................................................................................................5-18

5.6.1 Species Richness...........................................................................................................5-18

5.6.2 Spawning .......................................................................................................................5-18

5.6.3 Settlement and Early Associations with Habitat ............................................................5-18

5.6.4 Developmental Patterns and Cross-shelf Habitat Use ..................................................5-19

5.6.5 Trophic Patterns and Functional Groups .......................................................................5-20

5.6.6 Latitudinal Variations in Shallow Hardbottom Fish Assemblages..................................5-21
Table of Contents

(Continued)

Page 

Table of Contents iii 

CHAPTER 6 MARINE TURTLES..............................................................................................................6-1

6.1 INTRODUCTION...........................................................................................................................6-1

6.1.1 Adult Marine Turtles.........................................................................................................6-1

6.1.2 Hatchling Marine Turtles ..................................................................................................6-2

6.1.3 Predators..........................................................................................................................6-3

6.1.4 Water Depth.....................................................................................................................6-5

6.1.5 Seasonality.......................................................................................................................6-5

6.2 SPECIES OVERVIEWS................................................................................................................6-5

6.2.1 Green Turtles ...................................................................................................................6-8

6.2.2 Loggerhead Turtles..........................................................................................................6-9

6.2.3 Hawksbill Turtles............................................................................................................6-12

6.3 DISTURBANCE EVENTS AND TURTLES.................................................................................6-13

6.3.1 Shoreline Project Disturbances......................................................................................6-13

6.3.2 Red Tide Disturbances ..................................................................................................6-13

6.4 SUMMARY..................................................................................................................................6-13

Section 3 Functional Attributes of Nearshore Hardbottom and 

Mitigation Alternatives 

CHAPTER 7 FUNCTIONAL ECOLOGICAL ATTRIBUTES OF SHALLOW HARDBOTTOM.................7-1

7.1 INTRODUCTION...........................................................................................................................7-1

7.2 ECOLOGICAL FUNCTIONAL GROUPING..................................................................................7-2

7.2.1 Structure-Shelter Associated Functional Groupings........................................................7-3

7.2.2 Trophic Functional Groupings..........................................................................................7-9

7.3 LATITUDINAL COMPARISONS .................................................................................................7-13

7.4 DISTURBANCE ECOLOGY .......................................................................................................7-14

7.5 OBLIGATE AND FACULTATIVE HABITAT USE .......................................................................7-15

7.6 POPULATION CONNECTIVITY AND MAINTENANCE.............................................................7-17

CHAPTER 8 MITIGATING SHALLOW HARDBOTTOM IMPACTS.........................................................8-1

8.1 STRESSORS AND EFFECTS......................................................................................................8-1

8.1.1 Ecological Assessment Frameworks ...............................................................................8-1

8.1.2 Stressors, Responses, and Effects..................................................................................8-1

8.2 MITIGATION OF NEARSHORE HARDBOTTOM BURIAL ..........................................................8-9

8.2.1 Artificial Reefs ................................................................................................................8-10

8.2.2 Monitoring ......................................................................................................................8-15

8.3 CONCLUSIONS..........................................................................................................................8-17

8.3.1 Regional and Organism-specific Issues.........................................................................8-17

8.3.2 Monitoring Issues...........................................................................................................8-18
Table of Contents

(Continued)

Page 

Table of Contents iv 

CHAPTER 9 SUMMARY ...........................................................................................................................9-1

9.1 ALGAL, INVERTEBRATE, FISH, AND TURTLE ASSEMBLAGES..............................................9-1

9.1.1 Algae ................................................................................................................................9-1

9.1.2 Invertebrates ....................................................................................................................9-2

9.1.3 Fishes...............................................................................................................................9-3

9.1.4 Turtles ..............................................................................................................................9-4

9.2 ECOLOGICAL FUNCTIONS AND CONNECTIVITY ....................................................................9-5

9.3 MITIGATION .................................................................................................................................9-6 

9.4 RESEARCH NEEDS.....................................................................................................................9-6 

CHAPTER 10 LITERATURE CITED .......................................................................................................10-1

APPENDICES 

Appendix A....................................................................................................................................A-1 

Appendix B....................................................................................................................................B-1 

Appendix C................................................................................................................................... C-1 

Appendix D................................................................................................................................... D-1 
List of Tables v 

List of Tables

Table Page 

 1.1 Ecological terms used in the present study............................................................................1-7 

 2.1 Study counties from north to south with shallow hardbottom acreage by depth strata 

by year....................................................................................................................................2-1 

 2.2 Geological and biological comparisons between nearshore areas of the mainland 

east Florida coast and the northern margin of the Florida reef tract and the Florida 

Keys (Adapted from: Cooke, 1945; Kirtley and Tanner, 1968; Hoffmeister, 1974)................2-5 

 3.1 Macroalgal functional form group descriptions as proposed in Littler et al. (1983)................3-3 

 3.2 Effects of environmental conditions on the various algal functional form groups based 

on Littler et al. (1983)..............................................................................................................3-3 

 4.1 Species recruitment into intertidal and subtidal habitats from June 1997 through 

January 2000 in Boynton Beach, Florida ...............................................................................4-7 

 4.2 Abundance of associated macroinvertebrates collected from sabellariid worm reef 

(Stations 9 and 10 combined) at the St. Lucie Power Plant from April 1976 through 

April 1979 (From: Applied Biology, Inc., 1979).....................................................................4-15 

 4.3 Molluscs recorded during a survey of a newly constructed artificial reef structure in 

the nearshore (~8 m depth) area off Boca Raton, Florida (Adapted from: Cummings, 

1994).....................................................................................................................................4-16 

 4.4 Data summary for molluscs collected from 41 colonies of the scleractinian coral 

Oculina varicosa on four reefs off the east Florida coast (From: Reed and Mikkelsen, 

1987).....................................................................................................................................4-16 

 4.5 Comparison of worm reef arthropods found in the vicinity of Hutchinson Island, 

Florida with anthropod species found offshore in the same area (From: Camp et al., 

1977).....................................................................................................................................4-20 

 4.6 Abundance of associated decapods collected from sabellariid worm reef (Stations 9 

and 10 combined) at the St. Lucie Power Plant from April 1976 through April 1979 

(From: Applied Biology, Inc., 1979) ......................................................................................4-21 

 4.7 Comparison of associated decapod and stomatopod crustaceans from studies at 

Walton Rocks and the St. Lucie Power Plant (From: Applied Biology, Inc., 1979)..............4-21 

 4.8 The 51 most recurrent species of decapod and stomatopod crustaceans encountered 

from sabellariid worm reefs in the Indian River region of Florida, 1974 to 1975 (From: 

Gore et al., 1978)..................................................................................................................4-22 

 4.9 Species richness of decapod and stomatopod crustaceans collected from sabellariid 

worm reefs in the Indian River region of Florida, 1974 to 1975 (From: Gore et al., 

1978).....................................................................................................................................4-23 

 4.10 Rank analysis of intertidal decapods from the east Florida coast (From: van 

Montfrans, 1981) ..................................................................................................................4-23 

 4.11 Rank analysis of the dominant subtidal decapods from the east Florida coast 

(From: van Montfrans, 1981)................................................................................................4-24

 4.12 Total number of the three most common subtidal decapod species at stations along 

the east Florida coast (Adapted from: van Montfrans, 1981)...............................................4-24 
List of Tables

(Continued)

Table Page 

List of Tables vi 

 4.13 Numerical abundance of decapod species at Oculina varicosa coral reef stations at 

depths of 6, 27, 42, and 80 m (Adapted from: Reed et al., 1982)........................................4-25 

 4.14 Percent occurrence of decapod species at Oculina varicosa coral reef stations at 

depths of 6, 27, 42, and 80 m (Adapted from: Reed et al., 1982)........................................4-25 

 4.15 Physical parameters measured between 1976 and 1979 at Oculina varicosa coral 

reef stations at depths of 6, 27, 42, and 80 m......................................................................4-26 

 4.16 Abundance of isopods and amphipods from sabellariid worm rock with percentage of 

contribution of species to total abundance of each order at Sebastian Inlet, Florida, 

April 1984-March 1985 (From: Nelson and Demetriades, 1992) .........................................4-27 

 4.17 Summary of the life history characteristics of the ontogenetic stages of spiny lobster 

(Panulirus argus) (Adapted from: http://bio.fsu.edu/~herrnlab/Life_History_Table.pdf).......4-29 

 5.1 Focal groups of fishes based on trophic and habitat use attributes.......................................5-2 

 6.1 Role (functions) of nearshore hardbottom (NHB) (and reference to offshore 

hardbottom [OHB]) in marine turtle life history stages ...........................................................6-4 

 6.2 Marine turtle mark and recapture population studies conducted over nearshore 

hardbottom..............................................................................................................................6-7 

 6.3 Catch-per-unit-effort (CPUE) for marine turtle population studies conducted with 

tangle net capture methods adjacent to nearshore hardbottom habitats on the east 

coast of Florida .......................................................................................................................6-7 

 6.4 Macroalgal species present in foraging samples from various population studies 

conducted on marine turtles on nearshore hardbottom habitat............................................6-10 

 7.1 Total numbers of family, genera, and species of macroalgae associated with 

nearshore hardbottom habitat from studies along the east Florida coast ..............................7-2 

 7.2 Macroalgal species found on Florida’s Atlantic nearshore hardbottom habitat with 

known secondary metabolites proposed to aid in defense against herbivores....................7-16 

 8.1 Representative response categories in early stages of marine invertebrates or fishes 

when challenged by stressors from habitat burial (Modified from: Eisler, 1974; Olla et 

al., 1974).................................................................................................................................8-2 

 8.2 Invertebrate predictors of responses to dredge and fill burial: 1) organismal 

responses, and 2) assemblage-scale responses...................................................................8-5 

 8.3 Predictors of organism-scale responses to dredge and fill burial in fishes. ...........................8-7 

 8.4 Predictors of organism- and population-scale responses to dredge and fill burial in 

macroalgae.............................................................................................................................8-8 

 8.5 Predictors of organism- and population-scale responses to dredge and fill burial for 

marine turtles..........................................................................................................................8-9 

 8.6 Summary of compensatory mitigation for completed projects in southeast Florida 

between 1985 and 2004 (Adapted from: U.S. Fish and Wildlife Service, 2004)..................8-14 
List of Figures vii 

List of Figures

Figure Page 

 1.1 Overview of the seven-county project area............................................................................1-2 

 1.2 Nearshore hardbottom tide pools, Coral Cove County Park, south Jupiter Island, 

northern Palm Beach County, Florida ....................................................................................1-3 

 1.3 Broken pavement structure, nearshore hardbottom, central Brevard County, Florida...........1-4 

 1.4 Microhabitat complexity among diverse taxa under a nearshore hardbottom ledge, 

MacArthur Beach State Park, Palm Beach County, Florida...................................................1-4 

 1.5 Juvenile stage of the green sea turtle (Chelonia mydas) feeding on nearshore 

hardbottom algae, Boca Raton, Palm Beach County, Florida................................................1-5 

 1.6 Anglers fishing from nearshore hardbottom to capture pompano and other coastal 

pelagic species known to occur near sand-rock interfaces, central Brevard County, 

Florida...................................................................................................................................1-10 

 2.1 Aerial image of Bathtub Reef, Martin County, Florida............................................................2-3 

 2.2 Nearshore hardbottom algal and invertebrate colonization on ledge exposed at full 

moon low tide, Brevard County, Florida .................................................................................2-4 

 2.3 Aerial views of nearshore hardbottom structures and sediment change at northern 

Singer Island, Palm Beach County, Florida, 2003-2007 ........................................................2-7 

 2.4 Broward and northern Dade County plots from a summary of nearshore and 

offshore hardbottom information from the Southeast Area Monitoring and 

Assessment Program (SEAMAP) (Perkins et al., 1997) ........................................................2-8 

 2.5 Indian River County hardbottom from a summary of nearshore and offshore 

hardbottom (From: Perkins et al., 1997).................................................................................2-9 

 2.6 Florida Current (shown in orange) with offshore deflection near the southern 

boundary of Martin County, Florida (Image from: National Oceanic and Atmospheric 

Administration)......................................................................................................................2-10 

 2.7 Southward view of Bathtub Reef, Stuart, Martin County, Florida.........................................2-12 

 2.8 Paths of hurricanes and storms impacting east Florida during A) 2005 and B) 2004..........2-13 

 3.1 Path of the Florida Current (Gulf Stream) ..............................................................................3-4

 3.2 Percent composition of macroalgae and cyanobacteria divisions documented from 

studies conducted on nearshore hardbottom.........................................................................3-5 

 3.3 Percent composition of macroalgae and cyanobacteria functional form groups 

documented from studies conducted on nearshore hardbottom............................................3-6 

 3.4 Percent composition of macroalgal functional form groups (by county) documented 

from studies conducted on nearshore hardbottom.................................................................3-6 

 3.5 Representative species of the sheet functional form group growing on nearshore 

hardbottom: Ulva lactuca and Dictyota sp..............................................................................3-8 

 3.6 Percentage of species in the sheet functional group documented in studies 

conducted on nearshore hardbottom in counties from Brevard to Broward, Florida..............3-9 
List of Figures

(Continued)

Figure Page 

List of Figures viii 

 3.7 Percentage of species in the filamentous functional form group documented in 

studies conducted on nearshore hardbottom in counties from Brevard to Broward, 

Florida...................................................................................................................................3-10 

 3.8 Representative species in the coarsely-branched functional form group growing on 

nearshore hardbottom: Laurencia sp. and Caulerpa sp.......................................................3-11 

 3.9 Percentage of species in the coarsely-branched functional form group documented 

in studies conducted on nearshore hardbottom in counties from Brevard to Broward, 

Florida...................................................................................................................................3-11 

 3.10 Several representative species of the thick-leathery functional form group, including 

Agardhiella subulata and Gracilaria sp., growing on nearshore hardbottom .......................3-12 

 3.11 Percentage of species in the thick-leathery functional form group documented in 

studies conducted on nearshore hardbottom in counties from Brevard to Broward, 

Florida...................................................................................................................................3-13 

 3.12 Percentage of species in the jointed-calcareous functional form group documented 

in studies conducted on nearshore hardbottom in counties from Brevard to Broward, 

Florida...................................................................................................................................3-14 

 4.1 Intertidal worm reef at Bathtub Reef, Stuart, Martin County, Florida .....................................4-2 

 4.2 The trophic subweb postulated for the six most abundant species of decapod 

crustaceans on central Florida sabellariid worm reefs ...........................................................4-2 

 4.3 Close up of the anterior end of the reef-building polychaete Phragmatopoma 

lapidosa (caudata) ..................................................................................................................4-3 

 4.4 Close up of the anterior tube of an individual Phragmatopoma lapidosa (caudata) ..............4-3 

 4.5 Honeycomb structure created by thousands of Phragmatopoma lapidosa (caudata) ...........4-3 

 4.6 Monthly mean percentage cover of live adults, juveniles, destroyed worms, and 

vacant mounds for Phragmatopoma lapidosa on Bathtub Reef, Stuart, Martin 

County, Florida (McCarthy, 2008) ..........................................................................................4-4 

 4.7 Phragmatopoma lapidosa (caudata) recruits recovered after 5 weeks deployment in 

intertidal hardbottom off Boynton Beach, Florida ...................................................................4-5 

 4.8 High worm reef cover and wave activity at groynes at Palm Beach, Florida .........................4-5 

 4.9 Seasonal reproductive and life history trends of Phragmatopoma lapidosa in the 

Boynton Beach area as illustrated by McCarthy (2001).........................................................4-6 

 4.10 An intertidal colony of the scleractinian Siderastrea sp. in Palm Beach, Florida ...................4-8 

 4.11 The zooanthid Zoanthus pulchellus and various other sessile invertebrates (and 

green algae) on intertidal hardbottom in Palm Beach, Florida...............................................4-8 

 4.12 An intertidal colony of the rock-boring sponge Pione (Cliona) lampa in Stuart, Florida.......4-11 

 4.13 The demosponge Aplysina fistularia on hardbottom off Palm Beach, Florida .....................4-12 

 4.14 Colonial hydrozoans on a mitigation reef in Juno Beach, Florida ........................................4-12 

 4.15 Botryllus sp. overgrowing dead worm rock at Boynton Beach, Florida................................4-14 
List of Figures

(Continued)

Figure Page 

List of Figures ix 

 4.16 The volcano barnacle Tetraclita squamosa stalactifera and various other sessile 

invertebrates on intertidal hardbottom in Palm Beach, Florida ............................................4-17 

 4.17 Tidal spray crab (Plagiusa depressa) in Palm Beach, Florida .............................................4-18 

 4.18 Species composition at each of four Oculina stations (From: Reed and 

Mikkelsen, 1987) ..................................................................................................................4-31 

4.19 The rock-boring urchin Echinometra lucunter on shallow hardbottom habitat in Vero 

Beach, Florida ......................................................................................................................4-34 

 4.20 The sea cucumber Holothuria grisea on shallow hardbottom habitat in Ambersands 

Beach, Florida ......................................................................................................................4-34 

 5.1 Multispecies school of juvenile fishes including four species and three families 

(grunts, wrasses, and damselfishes) in association with nearshore hardbottom 

structure..................................................................................................................................5-1 

 5.2 Early juvenile lane snapper (Lutjanus synagris) associating with algal-rock-sand 

interface, 3 to 4 m, John U. Lloyd State Park, Broward County, Florida................................5-3 

 5.3 Resting nurse shark, Ginglymostoma cirratum, a generalized feeder on bottom 

invertebrates, 3 m depth, Palm Beach, Florida ......................................................................5-4 

 5.4 Newly settled grunts in the transitional epibenthic plantivore stage (approximately 

12 to 20 mm standard length) after settlement and before adoption of demersal 

feeding (by the late early juvenile stage), Palm Beach, Palm Beach County, Florida ...........5-6 

 5.5 Early life stages of primary reef-associating genera of haemulids and lutjanids of the 

western Atlantic ......................................................................................................................5-6 

 5.6 Comparisons of abundances of life history stages at two sites in northern Palm 

Beach County (From: Lindeman and Snyder, 1999)..............................................................5-7 

 5.7 Spawning coloration in male hairy blenny (Labrisomus nuchipinnis), with female in 

background, 2 m depth, Pecks Lake Reef, southern Martin County, Florida.......................5-10 

 5.8 A cryptic hardbottom resident, spotted moray eel (Gymnothorax moringa), depth 

2 m, MacArthur Beach State Park, Palm Beach County, Florida.........................................5-10 

 5.9 Example of nest of the sergeant major (Abudefduf saxatilis) on hardbottom, 20 m 

depth, Jupiter, Palm Beach County, Florida.........................................................................5-15 

 5.10 Early juvenile stages of reef herbivores including surgeonfishes, damselfishes, and 

parrotfishes on nearshore hardbottom, 3 m depth, Palm Beach, Palm Beach 

County, Florida .....................................................................................................................5-17 

 5.11 Relative adult trophic composition as measured by numbers of species per 

functional grouping ...............................................................................................................5-20 

 6.1 Adult female green turtles (in foreground and distance) crossing the nearshore 

hardbottom structure after nesting in Coral Cove Park, Palm Beach County, Florida...........6-1 

 6.2 Conceptual model of ontogenetic habitat stages in marine turtles (From: Musick and 

Limpus, 1997).........................................................................................................................6-2 
List of Figures

(Continued)

Figure Page 

List of Figures x 

 6.3 North Atlantic Gyre with potential oceanic transport routes for juvenile marine turtles 

(From: Musick and Limpus, 1997)..........................................................................................6-3 

 6.4 General locations of study areas for marine turtles in coastal areas superimposed 

over distribution of reef-building sabellariid worms along mainland east Florida 

(Modified from: Zale and Merrifield, 1989) .............................................................................6-6 

 6.5 Loggerhead (Caretta caretta) resting on offshore hardbottom, Breakers Reef, Palm 

Beach County, Florida............................................................................................................6-8 

 6.6 Juvenile green turtle (Chelonia mydas) over nearshore hardbottom in Palm Beach 

County, Florida .......................................................................................................................6-8 

 6.7 Juvenile hawksbill turtle (Eretmochelys imbricata) foraging on a tunicate wedged 

between the rocks on Nursery Reef, Pompano Beach, Florida (6 m water depth)..............6-12 

 7.1 Seasonal larval recruitment of the sabellariid worm Phragmatopoma lapidosa in 

intertidal and subtidal nearshore hardbottom habitats off Boynton Beach, Palm 

Beach County, Florida............................................................................................................7-6 

 7.2 Newly settled drums (Pareques species) commonly associate with ledges and 

triangular cavities at the base of shallow hardbottom structures, such as in this 

photograph from 2 m depth, Singer Island, Palm Beach County, Florida..............................7-7 

 7.3 Food web among six common decapod crustaceans from worm reefs 

(From: Gore et al., 1978)......................................................................................................7-10 

 7.4 Gray triggerfish, Balistes capriscus, pursuing invertebrate prey, Breakers Reef, 

approximately 6 m depth, Palm Beach, Florida....................................................................7-11 

 7.5 A juvenile French angelfish (Pomacanthus paru) cleaning an adult gray triggerfish 

(Balistes capriscus) in 4 m depth off Palm Beach, Palm Beach County, Florida.................7-12 

 8.1 Three scenarios of cross-shelf turbidity and sedimentation effects (direct and 

indirect) for systems subjected to A) no dredging, B) some dredging, and 

C) semi-continuous dredging (e.g., north-central Jupiter Island) ...........................................8-4 

 8.2 Artificial mitigation reef and substantial algal and hydroid growth, 6 m depth, Juno 

Beach, Palm Beach County, Florida ....................................................................................8-11 

 8.3 Prototype mitigation reef modules made from articulated mat with integral coquina 

rock surface ..........................................................................................................................8-12 
List of Acronyms xi 

List of Acronyms

BACI before-after-control-impact 

CEM conceptual ecological model

CERA comparative ecological risk assessment 

CL carapace length 

CPUE catch-per-unit-effort 

EBM ecosystem-based management 

ECB East Coast Biologists, Inc. 

EFH Essential Fish Habitat 

ESA Endangered Species Act 

FDEP Florida Department of Environmental Protection 

FP fibropapillomatosis 

FWC Florida Fish and Wildlife Conservation Commission 

FWRI Florida Wildlife Research Institute 

HAPC Habitat Areas of Particular Concern 

IHB intermediate hardbottom 

MHW mean high water mark 

NEPA National Environmental Policy Act 

NHB nearshore hardbottom 

NMFS National Marine Fisheries Service 

NOAA National Oceanic and Atmospheric Administration 

OHB offshore hardbottom 

PALM propagule and larval measurement 

SAB South Atlantic Bight 

SAFMC South Atlantic Fishery Management Council 

SCL straight carapace length 

SFCREMP Southeast Florida Coral Reef Evaluation and Monitoring Project 

SL standard length 

TL total length 

UMAM Uniform Mitigation Assessment Method 

USACE U.S. Army Corps of Engineers 

USEPA U.S. Environmental Protection Agency 

USFWS U.S Fish and Wildlife Service 
Section 1 

Project Objectives and Nearshore Hardbottom 

within the East Florida Seascape 

Section 1 

Project Objectives and Nearshore Hardbottom 

within the East Florida Seascape 
Section 1 1-1 

Project Objectives and Nearshore Hardbottom 

within the East Florida Seascape 

Chapter 1

Introduction

1.1 INTRODUCTION AND OBJECTIVES 

Nearshore hardbottom (NHB) habitats take many forms around the Greater Caribbean, from the coquina 

and worm reefs of mainland east Florida to the ironshore of the Bahamas and Turks and Caicos to the 

razor sharp dientes del perro (dog teeth) of northern Cuba (including much of Havana’s shore). There is 

a deservedly large research literature on a dozen or more geological or biological aspects of the deeper 

coral reefs of the Florida Keys; yet, there is a paucity of research information in peer-review journals on 

the approximately 400-km stretch of discontinuous, nearshore ahermatypic reefs of mainland east 

Florida, at the current northern limits of the subtropical Northwest Atlantic. 

Major coastal economies including sun-and-sand tourism and resident-driven recreational-use markets 

converge in mainland east Florida, with subtropical to warm temperate beaches intergrading with NHB in 

some locations between Miami/Dade County and Cape Canaveral (Figure 1.1). The beaches of this 

region are subject to many challenges to long-term sustainable management. Many of these challenges 

are amplified by continuing pressures for coastal development, diminishing natural resource status, and 

the probabilities of sea level rise. This distance is subject to semi-continuous wind and wave events and 

periodic erosion events. Southeast Florida is now a pending coastal mega-city. With a continued push of 

coastal growth northward, over 5 million people will soon exist within a narrow low-elevation corridor 

between Miami-Dade and Palm Beach Counties (Wallis et al., 2000). 

NHB habitats are the primary natural habitat at depths of 0 to 4 m for many organisms in this region. 

Currently, the majority of these structures are within a narrow, 200- to 400-m strip of the most nearshore 

marine waters and display a variety of structural forms, ranging from flat expanse with little relief to 

vertical mounds emergent at low tide to deeper structures that are much less subject to tidal effects. 

These habitats are patchily distributed from north to south, composing less than 50% of the alongshore 

distance of six of the seven county shorelines in the region and are often separated by kilometers of sand 

expanses. Indian River County is an exception, with NHB present along well over 50% of its shore, often 

extending into depths beyond 4 m (V. Kosmynin, Bureau of Coastal Systems, Florida Department of 

Environmental Protection [FDEP], Tallahassee, FL, personal communication). Structural complexity of 

the rock structures can vary latitudinally (Figures 1.2 and 1.3) and is enhanced by framework-building 

organisms such as tube-building polychaete worms (Gram, 1965; Kirtley and Tanner, 1968; Pandolfi et 

al., 1998; McCarthy, 2001), other invertebrates (e.g., sponges, anthozoans, bryozoans), and macroalgae 

(Goldberg, 1973; Gore et al., 1978; Nelson, 1989; Nelson and Demetriades, 1992). 

Situated among broad expanses of bare sand bottom, NHB structural features potentially serve a variety 

of ecosystem functions, including settlement and nursery areas, spawning sites, feeding areas, and 

shelter for hundreds of species of resident crabs, worms, shrimp, fishes, and many other animals and 

plants, many of which occur in close proximity (Figure 1.4). These functions translate into important 

ecosystem services such as improved water quality, recreational opportunities (e.g., fishing, surfing, 

diving for local communities), aesthetics, and wildlife habitat (Figure 1.5). 
Section 1 1-2 

Project Objectives and Nearshore Hardbottom 

within the East Florida Seascape 

Figure 1.1. Overview of the seven-county project area. Shallow hardbottom distributional information by 

county is presented in Table 2.1. 
Section 1 1-3 

Project Objectives and Nearshore Hardbottom 

within the East Florida Seascape 

Figure 1.2. Nearshore hardbottom tide pools, Coral Cove County Park, south Jupiter Island, northern 

Palm Beach County, Florida. Photo by D. Snyder. 
Section 1 1-4 

Project Objectives and Nearshore Hardbottom 

within the East Florida Seascape 

Figure 1.3. Broken pavement structure, nearshore hardbottom, central Brevard County, Florida. Photo by 

K. Lindeman.

Figure 1.4. Microhabitat complexity among diverse taxa under a nearshore hardbottom ledge, MacArthur 

Beach State Park, Palm Beach County, Florida. Sponges, tunicates, hydrozoans, 

macroalgae, and fish are present. Photo by D. Snyder. 
Section 1 1-5 

Project Objectives and Nearshore Hardbottom 

within the East Florida Seascape 

Figure 1.5. Juvenile stage of the green sea turtle (Chelonia mydas) feeding on nearshore hardbottom 

algae, Boca Raton, Palm Beach County, Florida. Photo by K. Jones. 

Based on consensus federal and state concerns over the declining condition of fisheries, reefs, and other 

coastal resources, some management policies have migrated towards potentially more precautionary, 

ecosystem-based management (EBM) tools, ranging from marine protected areas to increased protection 

of high value fishery habitats to interdisciplinary use of decision support tools to build more logical, 

multi-stakeholder governance policies (U.S. Executive Order on Coral Reefs, 1998; National Academy of 

Sciences, 1999; U.S. Commission on Ocean Policy, 2004). An ongoing example of the focus on new 

management approaches involves the recent listing of the elkhorn and staghorn corals (Acropora spp.) as 

threatened under the Endangered Species Act (ESA). Listing a colonial invertebrate species that also 

plays a large role for many other species as a structural habitat feature as threatened is an 

unprecedented step for ESA. This action has important implications for the management of NHB 

because the proposed critical habitat designation for ESA includes features essential to the conservation 

of threatened corals, i.e., natural consolidated hardbottom or dead coral skeleton that is free from fleshy 

macroalgae cover and sediment cover (South Atlantic Fishery Management Council [SAFMC], in prep.), 

to maximize the potential for successful recruitment and population growth. The recognition and 

designation of federal Essential Fish Habitat (EFH) is an earlier and broader example of the mandate for 

more application of EBM tools. NHB of the federal SAFMC’s four-state jurisdictional area (east Florida 

through North Carolina) was administratively designated as EFH in 1998 in response to a statutory 

mandate from the reauthorization (the Sustainable Fisheries Act of 1996) of the primary federal fisheries 

statute (the Magnuson-Stevens Fishery Conservation and Management Act of 1976). Federal action 

agencies that fund, permit, or carry out activities that may adversely affect EFH are required to consult 

with the National Marine Fisheries Service (NMFS) regarding the potential impacts of their actions on 

EFH, including consultations for coastal construction activities permitted or funded by the U.S. Army 

Corps of Engineers (USACE). The final rule on EFH determinations recognized that subunits of EFH may 

be of higher value. Such areas, termed Essential Fish Habitat-Habitat Areas of Particular Concern 

(EFH-HAPCs) by the SAFMC, can be identified using four criteria from the rule: a) importance of 

ecological functions; b) sensitivity to human degradation; c) probability and extent of effects from 

development activities; and d) rarity of the habitat. All NHB under SAFMC jurisdiction is also designated 
Section 1 1-6 

Project Objectives and Nearshore Hardbottom 

within the East Florida Seascape 

as EFH-HAPC (SAFMC, 1998); this potentially affects some life stages of the majority of the 73 species of 

10 families within the SAFMC Snapper-Grouper Management Plan (Lindeman et al., 2000). NHB is also 

designated as EFH in the SAFMC’s Fishery Management Plan for corals and coral reefs (SAFMC, 1982). 

The FDEP is central in efforts to minimize and mitigate impacts to NHB under Florida state administrative 

rules defining coastal construction permitting procedures. In east Florida, the primary source of 

anthropogenic impacts to NHB are large beach renourishment projects that involve the offshore dredging 

and inshore filling of up to 2,300,000 cubic meters of sediments per current project and can potentially 

result in effects upon various resources and habitats across the shelf (Marszalek et al., 1977; Nelson, 

1989; Bush et al., 2004). Increasingly important and expensive issues involve the use of mitigation 

projects to offset impacts, typically a key step in the finalization of permitting processes. It is in the best 

interest of all parties to characterize the functional attributes of NHB and apply that information to the 

evaluation of mitigation alternatives and final decision-making. To this end, the FDEP Bureau of Beaches 

and Coastal Systems has contracted the production of this report to synthesize available information 

pertaining to the ecological characteristics and functions of NHB habitat to help guide future management 

of these resources. 

Although many gaps are present, much information can be brought to bear on NHB ecological functions 

from an array of research, industry, and permitting literature that includes fields such as organismal and 

population biology, community ecology, coastal geology, physical oceanography, fishery science, 

engineering design, and economic valuation. Given the consensus need for EBM, the still complex policy 

questions involving hardbottom burial and mitigation, and the large but unsynthesized literature that 

already exists, this project uses existing and new information to address the following issues: 

1. What are specific ecological functions of NHB? 

2. What can happen to the coastal ecosystem (broader than NHB) when NHB is lost due to 

nourishment? 

3. Is the offshore hardbottom (OHB) habitat able to ecologically compensate for the loss of NHB? 

4. If the NHB habitat has specific properties and functions that cannot be compensated for by existing 

OHB habitat, what are the best mitigation alternatives for the loss of natural NHB? 

To address these issues, we assembled and examined relevant peer-reviewed journals and texts as well 

as unpublished or gray literature into a document hierarchically examining the primary assemblages, 

which collectively contain over 1,000 species. In part to manage this diversity, this synthesis report 

consists of three sections. Section 1 (Chapters 1 and 2) contains an introduction to the objectives, a 

physical description of the hardbottom, and terminology of this document. Section 2 (Chapters 3 to 6) 

contains assemblage-scale chapters on macroalgae and cyanobacteria, invertebrates, fishes, and marine 

turtles, reviewing known NHB diversity and functional groupings. Section 3 (Chapters 7 to 9) 

synthesizes the information from prior chapters to address functional ecology, stress characterization, 

and mitigation alternatives. The following sections of this chapter characterize NHB and OHB and 

examine fundamental ecological concepts and terms as they relate to this synthesis. 

1.2 ECOLOGICAL CONCEPTS AND TERMS 

Unlike the physical sciences, where seemingly immutable laws govern processes, ecology 

(Shrader-Frechette and McCoy, 1993; Sagoff, 2003) and evolutionary biology (Mayr, 1982, 2004) are 

founded on concepts. Unfortunately, there is much imprecision with usage and meaning of ecological 

terms and concepts. Foundational concepts such as diversity, stability, succession, community, and 

ecosystem can vary greatly in their meaning among ecologists. These problems are considerable, often 

commented on, and amplified by the difficulties in establishing what an ecosystem, community, or habitat 

actually is among different workers and perspectives (Strong et al., 1984; Underwood, 1986; Noss, 1996; 

Simberloff, 2004). 
Section 1 1-7 

Project Objectives and Nearshore Hardbottom 

within the East Florida Seascape 

The problem of defining communities was described by Whittaker (1975) as follows: “Most communities 

intergrade continuously along environmental gradients rather than forming distinct clearly defined zones.” 

A significant example of imprecise meaning was demonstrated by Grimm and Wissel (1997) for the term 

stability. They reviewed the ecological literature and found that the term stability had at least 

163 definitions based on 70 concepts. This is an unfortunate situation for managers or non-ecologists 

seeking scientific guidance. They concluded that a six-point checklist had major utility to “... classify 

ecological systems and provide a system of coordinates for communication” (Grimm and Wissel, 1997). 

To best manage ambiguity introduced by the imprecise terminology, basic ecological terms used 

throughout the text are defined in Table 1.1. We are not proposing these definitions for universal 

application, they simply have utility to the seascape and objectives addressed in this study. 

Table 1.1. Ecological terms used in the present study. Conceptual background and caveats summarized 

in text. 

Assemblage – a group of organisms defined according to phylogenetic, ecological, or other study-dependent criteria. 

Community – a group of species living and interacting together within a particular area.

Disturbance – a natural or anthropogenic perturbation that will disrupt the extant ecological processes of 

assemblages or successional processes within an area. 

Diversity – the number of species in a given area and their proportional abundances. 

Ecological functions – substantive ecological processes that occur within a community or ecosystem. 

Ecological services – the beneficial outcomes resulting from ecological functions that include human influence 

(e.g., fishing, snorkeling, bird watching, and nature photography). 

Ecosystem – an interacting group of biological communities and abiotic factors within a single geographic area. 

Ecosystem engineers – organisms that directly or indirectly modulate the availability of resources to other organisms 

by modifying, maintaining, or creating habitat architecture. 

Keystone species – particular species (or functional group) that influence control over critical processes that can 

affect the greater assemblage. 

Larval supply – the quantity or potential quantity of invertebrate or fish larvae that are transported into an area when 

competent to settle, metamorphose, and begin a benthic-associated existence. 

Metapopulation – a species system in which 1) local populations inhabit relatively discrete habitat patches and 

2) dispersal among patches allows demographic connectivity but local population dynamics remain relatively 

asynchronous from other populations. 

Opportunistic species – species with life history strategies that involve facultative, not obligate (dependent), resource 

use. Such species can utilize a wide array of shelter or food resources. 

Recruitment – a generic term referring to organismal entry into a given area. Specific examples include 

Larval recruitment – addition of new individuals into a population by arrival of incoming larvae into bottom 

habitats (e.g., larval settlement). 

Inter-habitat recruitment – shifts from one benthic habitat to another; not the first larval settlement event but a 

later habitat shift. 

Fishery recruitment – the size or age at which individuals are first retained by fishing gears (i.e., when they 

enter the fishery). 

Species richness – the total number of species within a project-specific area. 

1.3 NEARSHORE, INTERMEDIATE, AND OFFSHORE HARDBOTTOM 

For many comparative questions, distinguishing between NHB and OHB habitats is often complicated, 

particularly among differing latitudes in east Florida. Dozens of physical and biological continua co-vary 

within a highly dynamic shallow environment that does not obey strict rules about depth zonation. Many 

metrics therefore often respond in quasi- or non-linear manners to both natural and anthropogenic 

disturbances, prohibiting discrete line-drawing or flawless terminology. Nevertheless, means to 

distinguish measurement benchmarks (e.g., NHB and OHB) are logical and necessary to frame and 

inform answers to complex and interwoven questions about species, communities, fisheries, and coastal 

management. 
Section 1 1-8 

Project Objectives and Nearshore Hardbottom 

within the East Florida Seascape 

Our focus here is on comparative analyses of macroalgae, invertebrates, fishes, and marine turtles 

between NHB (0 to 4 m), intermediate hardbottom (IHB) (4 to 6 m), and OHB (>6 m) depths. The use of 

the 0 to 4 m depth range is based on the near-maximum depth of many NHB habitats and some 

associated biotic assemblages in the region. In addition, this can be a depth of the toe-of-fill subsequent 

to dredge-related burial of NHB. Within NHB, it can also be useful to refer to an intertidal area (0 to 1 m) 

and a subtidal area (1 to 4 m), which can be further subdivided in the case of some invertebrate and algal 

species. Among and within these intervals are gradual transitions in many population- and 

community-scale metrics, some of utility in examining ecological functions. We re-emphasize that the 

depth "boundaries" between NHB, IHB, and OHB are artificial benchmarks to allow some measureable 

stratification of a highly variable abiotic and biotic continuum; they are not self-contained zones. 

Currently, the majority of typically uncovered NHB structural features are within 200 m of the mean high 

water mark (MHW), though there are notable exceptions in which hardbottom structure is emergent 

west-to-east through two to three depth areas (NHB, IHB, and OHB), including Riomar Reef in St. Lucie 

County, Bathtub Reef and Peck’s Lake Reef in Martin County, Breakers Reef in Palm Beach County, and 

wide NHB eastern extensions in the area of Broward County (Fort Lauderdale). More detailed 

examination of NHB, IHB, and OHB occurrence and latitudinal variation is provided in Chapter 2. 

There can be substantial physical and biotic differences between NHB in the intertidal zone (0 to 1 m) and 

the subtidal zone (approximately 1 to 4 m). Both zones are populated by disturbance-mediated 

assemblages, and biotic distributional patterns can vary due to the dynamics of the physical environment. 

Disturbance examples include storms and the frequency and duration of hardbottom burial (more 

pronounced in fall and winter periods, and varying by latitude). The issue of ephemeral/seasonal 

changes in NHB is relevant to analyses of habitat functions discussed in Chapter 7. 

1.4 SPECIES: WINDOWS INTO ASSEMBLAGES? 

The use of particularly prominent species to stratify the analysis and/or management of assemblage 

complexity has been attempted for many decades in theoretical and applied ecology, and focal or 

indicator species have been discussed in detail. Simberloff (1998) reviewed a variety of constructs that 

have evolved for species-enhanced EBM, often based on single-species management models. 

Unusually representative or analytically valuable species constructs included indicator, flagship, umbrella, 

and keystone species used as diverse surrogates or representatives of a much wider array of species 

from local assemblages at risk from environmental impacts. Indicator species were considered tenuous, 

often because of imprecise metrics of performance and unclear objectives (Simberloff, 1998), though 

various reviews and many studies still employ differing types of indicator species (Zacharias and Roff, 

2001). Umbrella species, those needing so much habitat for protection that other species are protected 

as well, could be useful, but are typically constrained in reflecting fundamental community functions and 

measurement, as are flagship species (Walpole and Leader-Williams, 2002). A possible umbrella species 

in NHB areas is the sabellariid reef-building worm Phragmatopoma lapidosa (Chapters 4 and 7). 

In part based on the work of Paine (1969) on nearshore rocky intertidal community changes, Simberloff 

(1998) concluded that keystone species played the most significant role in influencing assemblage 

structure and could potentially be useful tools for multispecies management. A fundamental criterion is 

that removal of the keystone species should substantially shift assemblage structure. In addition, the 

roles of some keystone species are highly context-dependent (Simberloff, 1998). Based on empirical 

information from Australian reefs, Bellwood (2008) has identified a phenomenon termed sleeping 

functional groups or species. In one case, a siganid rabbitfish of little prior significance in most metrics 

assumed a keystone-like role when the long-standing system of macro-herbivorous fishes changed in 

response to a fishing-induced phase shift. 

Zacharias and Roff (2001) concluded that the keystone species concept was both relatively strong and of 

varying utility in management. They also concluded that it was possible to define and measure indicator 

species under some conditions and suggested two categories: composition indicators and condition 

indicators. The former species reflect a specific niche or range of measurable environmental tolerances 

and are relatively independent of sample size. Condition indicators are species that somehow “provide 
Section 1 1-9 

Project Objectives and Nearshore Hardbottom 

within the East Florida Seascape 

an assessment over a range of stress” and can “differentiate between natural and anthropogenic stress” 

(Zacharias and Roff, 2001). With the exception of keystone species, we do not currently use any of these 

other terms. However, some NHB species may have utility as indicator species in some studies 

(e.g., black margate, Anisotremus surinamensis, as a composition indicator), if specific indicator criteria 

are well-defined. 

1.5 EQUILIBRIUM AND SCALE 

In addition to ambiguity in concepts and terms, agreement between theory and application in ecology has 

been generally lacking. Conceptual paradigms in ecology today may be broadly divided into equilibrium 

and non-equilibrium views (Wiens, 1984; Walter and Hengeveld, 2000; Rohde, 2005; Wallington et al., 

2005), and, as with most dichotomies, the truth probably lies somewhere in between the two poles. 

Theory formulated before the 1990’s was largely based on notions of equilibrium in natural assemblages 

(e.g., MacArthur, 1972). For example, this paradigm viewed ecological succession as a linear sequence 

of events leading to a stable climax, with any disturbance to that sequence considered something outside 

of the “system.” This viewpoint has been replaced by more probabilistic and non-equilibrium viewpoints 

(e.g., Botkin, 1990; Sale, 1991; Shrader-Frechette and McCoy, 1993; Drury, 1999; Rohde, 2005; 

Wallington et al., 2005). 

In the non-equilibrium world, disturbance is considered an inherent property of the ecosystem and 

equilibria are artifacts of observation, not real properties of the system. Unfortunately, many of these new 

ideas have not been translated into the regulatory arena (Wallington et al., 2005). Schrader-Frechette 

and McCoy (1993) summarized key conceptual issues and emphasized the importance of case study 

approaches, contending that ecological problem-solving would be most effective when ecological 

knowledge (natural history), not ecological theory, was applied. 

In this report, a basic conceptual assumption supported by the data is that biota inhabiting NHB are not in 

any kind of static or spatial equilibrium over long time scales. We state this now and throughout the text 

to ensure that managers realize the implications of a strict equilibrium stance (Chapters 7, 8, and 9) 

when assessing impacts and the results of mitigative measures employed to offset those impacts (Wiens 

and Parker, 1995; Parker and Wiens, 2005). 

Another fundamental component is that scale and assessment of equilibrium is inherently based on the 

scales of the system under examination. The structure of all levels of ecological hierarchies depends 

upon spatial and temporal scale. Consideration of spatial scale is paramount to an understanding of 

assemblage patch dynamics, particularly in disturbance-mediated environments such as NHB (Levin and 

Paine, 1974; Pickett et al., 1989; Wiens, 1989; Kotliar and Wiens, 1990; Wu and Loucks, 1995). The 

relationships between species richness and functional diversity (Hooper et al., 2002) can be tenuous but 

potentially useful in comparing NHB and OHB functional equivalencies and dissimilarities. 

1.6 ECOLOGICAL FUNCTIONS 

Several project objectives involve identifying ecological functions within natural and artificial habitat types 

and the subsequent comparison of functions among subregions. Function can be addressed in terms of 

processes and in terms of functional groups of characteristics and similar species (sometimes known as 

guilds) (Wilson, 1999). A major challenge at the interface of ecological research and applied 

management involves the difficulties in merging the conceptual landscape implied by the term function

with immediate political management needs. 

The conceptual underpinnings and terminology associated with the word function are highly variable, 

encompassing dozens of metrics. At least four broad meanings for the term function were identified by 

Jax (2005): 1) processes of changes of state (e.g., organismal feeding); 2) the merging of multiple 

processes in a whole system context (e.g., system functioning); 3) ecological roles within systems (e.g., 

functional groups such as producers or consumers); and 4) particular services of the system to humans 

and society (e.g., ecosystem services such as pollination and photosynthesis). Based on a 

comprehensive examination of the literature on NHB and ongoing examination of much of the literature 
Section 1 1-10 

Project Objectives and Nearshore Hardbottom 

within the East Florida Seascape 

on functions and functional groups, we identified nominal functional groupings within shallow hardbottom 

assemblages. Studies of habitats often have focused on two basic categories of functional attributes: 

1) structure and shelter resource use and 2) feeding assemblage relationships. We use the following 

non-exhaustive categories that are compatible with the framework employed by the Uniform Mitigation 

Assessment Method (UMAM) protocol (e.g., Continental Shelf Associates, Inc. et al., 2006). 

• Structure-shelter associated functional groupings. Examples include groups of species sharing one 

or more of the following attributes and associations with NHB through OHB: 

− Habitat engineering functions (keystone framework builder); 

− Nesting and spawning site usage; 

− Substrate as attachment resource for sessile organisms; 

− Settlement and post-settlement nursery area usage; and 

− Juvenile and adult patterns of cross-shelf distribution. 

• Trophic functional groupings. Food capture patterns, larger energetic fluxes, and physical dynamics 

drive food web dynamics (e.g., intercommunity nutrient exchange functions of carnivores and 

herbivores on NHB and OHB [Parrish, 1989]), such as the following: 

− Herbivory; 

− Carnivory; 

− Autotrophy; 

− Detritivory and omnivory; and 

− Cleaning symbioses. 

The relationship between ecological function 

analysis and societal measures has a 

considerable history (Hooper et al., 2002). 

Human societies can describe functions within 

an ecosystem and functions of the whole system 

(Jax, 2005). Tens of thousands of year-round 

local residents use the beach not only for 

sunbathing but also to take advantage of the 

nearshore waters, occasional reefs, and many 

coastal recreational activities (Figure 1.6) 

available in the warmest waters of the 

continental U.S., which also offer some of the 

highest coastal biodiversity. The services 

provided by nearshore reefs for humans are 

often family-based and multi-generational, 

including fishing, diving, surfing, and other forms 

of recreational resource use. 

Figure 1.6. Anglers fishing from nearshore hardbottom 

to capture pompano and other coastal 

pelagic species known to occur near 

sand-rock interfaces, central Brevard 

County, Florida. Photo by K. Lindeman. 
Section 1 2-1 

Project Objectives and Nearshore Hardbottom 

within the East Florida Seascape 

Chapter 2

Shallow Hardbottom of East Florida and the

Regional Shelf Setting

2.1 INTRODUCTION 

The study region is bounded on the south by Government Cut, the southern boundary of Miami Beach, 

(Figure 1.1). To the north, the study region is bounded by Port Canaveral. There are geological, 

biological, and climatic differences between the northern and southern portions of the study region. For 

many coastal marine organisms, the zoogeographic break between subtropical and warm temperate 

regions typically occurs between the Jupiter Inlet and Cape Canaveral, approximately 230 km to the north 

(Briggs, 1974; Gilmore, 1995). Transitional warm temperate, wide-shelf systems compared to 

subtropical, narrow-shelf systems are increasingly represented north and south of the Jupiter Inlet. 

Seven of Florida’s 35 counties with marine shores are within the study region. Table 2.1 provides the 

shoreline lengths and NHB acreages per year available. Brevard and Palm Beach Counties have the 

longest shorelines within the full project area, while the shorelines of the remaining counties are highly 

similar in length, ranging from 34.5 to 38.4 km. Primary economic drivers are concentrated in the 

southern three counties and pushing economic growth and coastal construction into northern counties 

where relatively less-disturbed coastal reaches can be found. There are substantial structural and NHB 

areal variations present across the 355-km north-south project reach, as well as considerable variation in 

the availability of areal NHB data (Table 2.1). The information in Table 2.1 is further evaluated in 

Section 2.1.2 and later chapters. 

Table 2.1. Study counties from north to south with shallow hardbottom acreage by depth strata by year. 

Most data were provided from county agencies. 

County 

Latitude at 

Southern Border 

(N) 

Shoreline Length 

km (mi) 

% Shoreline in 

Study Region 

Total 

Hardbottom 

(acres) 

Locationc Year 

Brevard 27° 51 ́ 114.9 (71.4) 32 42.3 R-70 to R-118 2004 

Indian River 27° 33 ́ 36.1 (22.4) 10 278.5a R-37 to R-49 2007 

St. Lucie 27° 15 ́ 34.6 (21.5) 10 56.6 R-77 to R 118 2008 

Martin 26° 58 ́ 34.5 (21.4) 10 N/A N/A N/A 

27.0 R-59 to R-71 2007 

337.4 County 2004 

283.3 County 2003 

4.2 R-59 to R-71 2003 

361.3 County 2001 

375.0 County 2000 

8.8 R-59 to R-71 2000 

16.8 R-59 to R-71 1998 

Palm Beach 26° 19 ́ 72.8 (45.2) 21 

444.1b

 County 1993 

Broward 25° 58 ́ 38.4 (23.9) 11 125.2 County N/A 

Miami-Dade 25° 46 ́ 23.6 (14.7)d 7 N/A N/A N/A 

Total Shoreline Length --- 354.9 (220.5) 100% --- --- 

a

 Mapped out only to ~450 m offshore; hardbottom beyond this is not quantified. 

b

 Includes areas within inlets that were not counted in other years. 

c

 Florida Department of Environmental Protection Reference Monument. 

d

 Measured to southernmost point of Miami Beach (north side of Government Cut). 

N/A = Not available. 
Section 1 2-2 

Project Objectives and Nearshore Hardbottom 

within the East Florida Seascape 

2.2 GEOLOGICAL AND BIOLOGICAL ATTRIBUTES OF NEARSHORE HARDBOTTOM 

Most NHB structures of mainland east Florida are current marine components of the Anastasia formation, 

which is composed of sediments and mollusc shells (particularly the coquina clam Donax). This material 

accumulated on shorelines during the last Pleistocene interglacial period approximately 80,000 to 

120,000 years ago and was later lithified during low sea level stands (Cooke and Mossom, 1929; Cooke, 

1945; Duane and Meisburger, 1969). The bulk of the NHB along mainland east Florida is composed 

primarily of this lithified shellrock; however, there is a transition in the area of Deerfield Beach in north 

Broward County where the Anastasia formation intergrades with and may be replaced to the south by the 

Miami Oolite formation and other features (Cooke, 1945; Hoffmeister, 1974). These limestone strata are 

significant features with east-west reaches that extend across coastal east Florida and still represent 

terrestrial landscape features. Prominent coastal ridges of these features occur along U.S. Highway 1 

throughout the study region (e.g., Rockledge, Brevard County and Silver Ridge, Coconut Grove, 

Miami-Dade County). 

NHB habitats are the natural ahermatypic reef structures at depths of 0 to 4 m of this region, and a 

diverse array of users refer to them by names that include coquina reefs, worm rock, Anastasia outcrops, 

worm reef, and nearshore reefs. In some areas, NHB reaches heights of 1.5 to 2 m above the bottom 

and is highly convoluted (Figure 1.2). In other areas, NHB is low-relief and forms pavement-like 

surfaces, often with small overhangs and crevices (Figure 1.3). Currently, little information is readily 

available on in-situ differences in physical and biotic properties of submerged Anastasia NHB compared 

to submerged NHB of differing origin in Broward County. 

NHB habitats are patchily distributed among expanses of relatively barren sediments (Table 2.1; 

Figure 2.1) and commonly possess worm reef structure in shallow areas from southern Palm Beach 

County to the north. NHB habitats of the region show very low coral diversities, yet have moderate to 

high diversities of other invertebrates, algae, and fishes. Nelson (1989) recorded 325 species of 

invertebrates and plants from NHB habitats at Sebastian Inlet. Hard corals are rare or uncommon in 

many NHB patches due to high turbidities and wave energy, but several species can be locally present 

(e.g., Oculina diffusa and O. varicosa in St. Lucie County; Acropora cervicornis in Broward County). Both 

Siderastrea radians and S. siderea can occur on shallow hardbottom in all counties from Brevard to 

Dade. A locally prominent contributor to habitat structure and biological diversity of NHB along east 

Florida is the polychaete Phragmatopoma lapidosa, also known as P. caudata (Nelson and Demetriades, 

1992; Kirtley, 1994; Drake et al., 2007). Worms of this species (Family Sabellariidae) settle on 

hardbottom and glue together sedimentary particles of specific sizes and origins to build sand tubes, 

forming shallow reefs in intertidal and shallow subtidal hardbottom areas (Gram, 1965; Kirtley and 

Tanner, 1968; Pandolfi et al., 1998; McCarthy, 2001). This species meets all the attributes and is the 

primary habitat engineer (Jones et al., 1994; Coleman and Williams, 2003; Jones et al., 2004) of NHB 

communities in mainland east Florida. Its distribution extends southward to Santa Catarina, Brazil 

(Kirtley, 1994), though it is often uncommon or absent on many islands and the Florida Keys (McCarthy et 

al., 2008). There are significant differences in the invertebrate fauna of intertidal NHB habitats in 

comparison to subtidal NHB habitats, driven in part by the high abundance of P. lapidosa in intertidal NHB 

habitats (McCarthy et al., 2003; also see Chapters 4, 5, and 7). 

In east Florida, the structure provided by NHB and associated habitat engineers (e.g., polychaete worms) 

supports a higher diversity and abundance of many marine invertebrate species than that of neighboring 

sand or hardbottom habitats (Gore et al., 1978; Nelson, 1989). The differences between areas with and 

without hardbottom can exceed an order of magnitude in terms of both species richness and abundance. 

At least 8 federal- and 15 state-listed species are associated with nearshore reefs off east Florida 

(U.S. Fish and Wildlife Service [USFWS], 1999), and NHB is considered an important source of food and 

shelter for juvenile green turtles (Chelonia mydas) (Ehrhart et al., 1996; Wershoven and Wershoven, 

1989; Holloway-Adkins, 2001). 
Section 1 2-3 

Project Objectives and Nearshore Hardbottom 

within the East Florida Seascape 

Figure 2.1. Aerial image of Bathtub Reef, Stuart, Martin County, Florida. 
Section 1 2-4 

Project Objectives and Nearshore Hardbottom 

within the East Florida Seascape 

Prior to the current synthesis report, NHB was recorded to provide shelter for over 325 invertebrate 

species (Gore et al., 1978; Nelson, 1988, 1989; Nelson and Demetriades, 1992) and 192 fish species 

(Gilmore, 1977; Gilmore et al., 1981; Lindeman, 1997a; Lindeman and Snyder, 1999). Substantial 

geological evidence suggests that NHB and worm reefs are also important in the maintenance and 

persistence of beaches and barrier islands by retention of sediments and the progradation of beaches 

(Kirtley, 1966, 1967; Multer and Milliman, 1967; Gram, 1965; Kirtley and Tanner, 1968; Mehta, 1973; 

Kirtley, 1974; Pandolfi et al., 1998). It is important to emphasize, however, that the vast majority of 

hardbottom is not encrusted by nor derived from tube worms (Figure 2.2). Worm reefs also can 

monopolize space, precluding other algae and sessile invertebrates. 

Figure 2.2. Nearshore hardbottom algal and invertebrate colonization on ledge exposed at full moon low 

tide, Brevard County, Florida. Algal species represented include Caulerpa racemosa, 

C. prolifera, Ulva lactuca, Laurencia sp., Aghardiella subulata, Gelidiopsis sp., and others. 

Cuban stone crabs (Menippe nodifrons) are visible in ledge on lower left. Photo by K. Lindeman.

NHB structures are largely carbonate composed via lithification of biogenic components (often calcareous 

microstructures of the small coquina bivalve Donax) and current framework-building organisms, 

particularly P. lapidosa (=caudata). Is the structure typical of NHB properly termed a reef? A 

fundamental way to examine this question is to go to the original source marine geological literature that 

described east Florida NHB. Early papers refer to NHB structures as reefs with geological and biological 

rationale, often including the role of the sabellariid worm species P. lapidosa in beach stabilization (Gram, 

1965; Kirtley, 1967; Kirtley and Tanner, 1968). These structures reflect and absorb wave energy, 

influence sediment movements, maintain structure position over time, support substantial biotic 

populations, and display fundamental attributes of ahermatypic, inorganic, or marginal reefs as described 

in Gram (1965), Kirtley and Tanner (1968), Fagerstrom (1987), Pandolfi et al. (1998), and Kleypas et al. 

(2001). Ichthyofaunal compositions are dominated by reef-associated species (Lindeman and Snyder, 

1999; Baron et al., 2004). 
Section 1 2-5 

Project Objectives and Nearshore Hardbottom 

within the East Florida Seascape 

The term NHB is also applied in the Florida Keys (Hoffmeister, 1974; Chiappone and Sullivan, 1996). 

NHB habitats of the Florida Keys can differ both geologically and biologically from mainland east Florida 

areas (Table 2.2). Emergent upland components of the Florida Keys are derived from ancient reefs of 

the Florida Reef Tract and typically do not have sizeable beaches nor a nearshore current regime for 

delivery of beach-quality sediments. Compared to mainland east Florida, most Florida Keys NHB habitats 

are distributed among areas with higher organic sediments, increased seagrasses, more corals, and 

reduced wave conditions, though there are exceptions offshore (Ginsburg, 1956). In contrast to the Keys, 

beach systems are common in mainland east Florida areas and their geologically distinct, sedimentary 

barrier islands (Table 2.2). 

Table 2.2. Geological and biological comparisons between nearshore areas of the mainland east Florida 

coast and the northern margin of the Florida reef tract and the Florida Keys (Adapted from: 

Cooke, 1945; Kirtley and Tanner, 1968; Hoffmeister, 1974). Transition areas are given for 

each attribute. 

Geological or Biological 

Component 

Mainland 

North of Transition 

Geographic 

Transition Areas 

Florida Keys 

South of Transition 

Island type Sedimentary barrier islands Key Biscayne – Soldier Key Coral/limestone islands 

Bedrock type Anastasia limestone Palm Beach – Broward Counties Miami Oolite and limestone 

Sabellariid worm reefs Locally common Broward – Dade Counties Rare 

Shallow corals Uncommon Key Biscayne – Soldier Key Common 

Seagrasses Absent Miami Beach – Fisher Island Abundant 

Predominant sediment type Quartz – calcium carbonate Key Biscayne – Soldier Key Calcium carbonate – quartz 

Predominant sediment size Coarse Key Biscayne – Soldier Key Fine 

Wave energy Intermediate to high Martin – Broward Counties Low 

2.3 DISTRIBUTION OF SHALLOW HARDBOTTOM IN EAST FLORIDA, MIAMI TO 

CANAVERAL 

Recent information on distributions and areal amounts of shallow hardbottom habitats are housed most 

comprehensively at the county scale from Miami to Canaveral (Dade through Brevard Counties, 

respectively) by the responsible county agencies and, at the federal level, by the USACE and NMFS in 

the form of various National Environmental Policy Act (NEPA)-associated documents from past dredge 

and fill projects. Table 2.1 summarizes some of the currently available macro-level information on NHB 

acreage and distributions. The construction of coastal inlets probably has contributed to the exposure of 

NHB in some areas by interrupting the littoral movement of sand. Based on available information 

(i.e., county mapping products), the counties with the most NHB are Palm Beach and Indian River, each 

appearing to have at least 250 acres of areal coverage. This value is essentially an average of a single 

snapshot (aerials) taken annually in each county. Broward has significant amounts of IHB at depths of 

4 to 6 m, with NHB as well. NHB is distributed near ridge complexes of substantial geomorphological 

complexity that occupy at least 125 acres (K. Banks [Broward County Department of Environmental 

Protection], B. Walker [National Coral Reef Institute, Nova Southeastern University Oceanographic 

Center], and Q. Robertson [Coastal Planning & Engineering, Inc.], pers. comm.). Martin and St. Lucie 

counties have comparatively less NHB, but can have substantial outcrops. Dade currently has no shallow 

NHB (<4 m water depth), though hardbottom with limited epibiota occurs in some areas around the 4 m 

depth contour, often grading into more densely colonized hardbottom by 6 m depths (B. Flynn, 

Miami-Dade Department of Environmental Resources Management, pers. comm.). Brevard’s NHB is 

very discrete, consisting of an approximately 40-acre system in the central part of the county that does 

not intergrade with deeper hardbottom. 

Table 2.1 is not comprehensive, as some data from aerial overflight imagery are not available (or have 

not been digitized to delineate hardbottom), and a full survey of other information sources (e.g., past 

NEPA documents) was outside the scope of this project. In addition to data summaries such as 

Table 2.1, aerial imagery can be assembled to show NHB distribution patterns among the seven counties 
Section 1 2-6 

Project Objectives and Nearshore Hardbottom 

within the East Florida Seascape 

for the years when mapping was conducted. In addition, there are substantial state and federal efforts 

underway both to map hardbottom resources as funding develops and also to gather existing information 

in publicly accessible manners (e.g., www.safmc.net). 

Evaluating NHB areal distributions and biological relationships is complicated by the burial dynamics of 

lower elevation NHB outcrops resulting from the many storms and wave events in the region (Nelson, 

1989; McCarthy et al., 2003; Bush et al., 2004; Continental Shelf Associates, Inc., 2006; Lybolt and Tate, 

2008; Montague, 2008). Therefore, the presence of hardbottom in mapping efforts is seasonally and 

annually variable according to trends in local wind and wave activity and extra-regional storms; such 

trends may often vary interannually as well. Imagery of the system shows substantial hardbottom 

structures and their point-in-time position on the seascape for multi-year time steps, as at Singer Island in 

Palm Beach County where acreage has increased approximately three-fold in recent years (Figure 2.3). 

Typically, more NHB is exposed during summer and lower overall nearshore energy periods (also the 

only feasible photographic periods for most areas). 

The South Atlantic Bottom Mapping Work Group of the Atlantic States Marine Fisheries Commission 

generated hardbottom mapping information for the South Atlantic Bight, including the study region from 

the beach to a depth of 200 m. The project ended in 2007, and the information is available at 

www.safmc.net. Figures 2.4 and 2.5 are from a summary of NHB and OHB mapping information from 

the Southeast Area Monitoring and Assessment Program (SEAMAP) (Perkins et al., 1997). There are 

fundamental differences in cross-shelf NHB habitat distribution among counties (e.g., Broward and 

northern Palm Beach). In Broward, the level of hardbottom structural overlap of the NHB, IHB, and OHB 

depth ranges exceeds that of many reaches to the north, with particular exceptions in southern Martin 

and Indian River Counties. 

Depth contours at 4 and 6 m are highly variable in terms of distance from shore throughout the study 

region. This complicates efforts to segregate patterns according to the nominal NHB, IHB, and OHB 

categories (Figure 2.1). The available data by county are also patchy in quantity and quality, and the 

ability to accurately segregate data by the three nominal categories currently is not possible for most 

counties. Nonetheless, there are many areas of Palm Beach and Brevard Counties, for example, where 

the NHB is not continuous with IHB and OHB. The NHB, IHB, and OHB designations are useful 

benchmarks for beginning to examine the complex interrelationships between the shallowest hardbottom 

assemblages and those in depths greater than 6 m. The challenges are amplified by the absence of 

depth-stratified information for most taxa and localities as well as by the dynamic spatial environment. 

2.4 OFFSHORE HARDBOTTOM AREAS 

Recognizing the difficulties in defining depth boundaries discussed earlier, this study uses a depth range 

of >6 m to represent OHB. There are many ways in which to characterize hardbottom and live coral 

habitats from >6 m to the shelf edge, with most available studies concentrated on the southern part of the 

project region (Goldberg, 1973; Moyer et al., 2003; Banks et al., 2008). We refer to shallow OHB 

(6 to 10 m) and midshelf OHB (>10 m) based on physical and biotic characteristics that are actually 

distributed across a continuum (see Chapter 1 regarding the high permeability of these depth “ranges”). 

In the present study, our emphasis is almost entirely on shallow OHB compared to midshelf OHB. 

Complex physical and biological assemblage relationships exist between OHB and the intermediate, 

transitional depth range for hardbottom, 4 to 6 m (IHB). OHB from shallow and midshelf areas potentially 

receives a portion of the grunt, wrasse, and other species of fish and invertebrate settlers that survive and 

eventually redistribute into deeper waters, sometimes across the broader shelf. These relationships are 

discussed by major biotic group in Chapters 3, 4, 5, and 6, as well as in Chapter 7 with respect to 

ecological functions. 

The relict Holocene reefs from Palm Beach to Dade Counties are the most-examined within the study 

region, with Lighty (1977), Moyer et al. (2003), and Banks et al. (2007) referring to shore-parallel inner, 

middle, and outer reefs. The latter papers also refer to a nearshore beach ridge complex (see 

Chapter 1); this is the depth zone of NHB. Though not frame-building, the hardbottom across the three 

reef lines is colonized by reef fauna characteristic of the Caribbean (Goldberg, 1973; Banks et al., 2008). 
Section 1 2-7 

Project Objectives and Nearshore Hardbottom 

within the East Florida Seascape 

Figure 2.3. Aerial views of nearshore hardbottom structures and sediment change at northern Singer 

Island, Palm Beach County, Florida, 2003-2007. 
Section 1 2-8 

Project Objectives and Nearshore Hardbottom 

within the East Florida Seascape 

Figure 2.4. Broward and northern Dade County plots from a summary of nearshore and offshore 

hardbottom information from the Southeast Area Monitoring and Assessment Program 

(SEAMAP) (Perkins et al., 1997). 
Section 1 2-9 

Project Objectives and Nearshore Hardbottom 

within the East Florida Seascape 

Figure 2.5. Indian River County hardbottom from a summary of nearshore and offshore hardbottom 

(From: Perkins et al., 1997). 
Section 1 2-10 

Project Objectives and Nearshore Hardbottom 

within the East Florida Seascape 

The inner reef crests at a depth of approximately 8 m and consists of discontinuous patch reef outcrops 

along a shore-parallel axis that periodically form elongate reefs (Banks et al., 2008), representing the 

OHB and IHB components of the southern project region in the present study. Throughout most of Palm 

Beach County, however, no inner or middle reefs are found (Banks et al., 2008). The nature and extent 

of shallow OHB varies with latitude along the study area. In some areas, most notably Breakers Reef, 

Palm Beach; Bathtub Reef and Peck’s Lake, Stuart; and Riomar, Vero Beach (Figure 2.5), hardbottom 

extends seaward from NHB to OHB depth ranges. In most other areas, NHB is not continuous with 

hardbottom in greater depths. 

Epibiota colonizing shallow OHB vary in taxonomic composition and diversity in both north-south and 

cross-shelf directions. Summaries for the Palm Beach through Dade County area are provided in Banks 

et al. (2008). Variations in light penetration, water temperature, sedimentation, and circulation are 

particularly present in shallow hardbottom areas and greatly influence the structure and dynamics of 

epibiotic assemblages (Rogers, 1990). Moyer et al. (2003) found substantial differences in benthic 

communities between the inner reef and the middle and outer reefs. For example, densities of octocorals 

and sponges were lower on the inner reef, presumably in response to greater environmental variability in 

shallower waters. There has been little directed study of hardbottom and reef structures and their 

epibiotic assemblages north of the Palm Beach area. General trends such as the north-south gradient in 

species diversity and basic taxonomic composition have been described peripherally for some epibiotic 

taxa, including corals and algae (Humm, 1969; Briggs, 1974; van den Hoek, 1975; Searles and 

Schneider, 1980; Jaap, 1984), but specific details of most assemblages in 6 to 10 m depths within the 

region remain poorly known. Taxa-specific discussions of known shallow OHB, IHB, and NHB 

associations among fishes, algae, invertebrates, and marine turtles are presented in Section 2 and 

summarized in terms of functional ecological attributes in Section 3. 

2.5 REGIONAL SETTING: THE FLORIDA CURRENT AND THE CONTINENTAL SHELF 

The continental shelf and associated waters off the southeastern U.S., collectively termed the South 

Atlantic Bight, extend from Cape Hatteras, North Carolina to southeast Florida. Shelf widths vary from 

just 3 km off Palm Beach County, Florida to a maximum of 120 km off 

Georgia. Climatically and biogeographically, the area contains the 

transition region between many tropical and warm temperate 

organisms of the Greater Caribbean and the Carolinas. Influences on 

oceanographic regimes of the continental shelf of mainland east 

Florida include 1) activities of the Florida Current and meanders of the 

western frontal boundary; 2) seasonal heating and cooling; and 

3) bottom topography. Winds and tides can also modify circulation 

patterns, especially nearshore or where density gradients are weak 

(SAFMC, in prep.). Relatively brief consideration of the underlying 

oceanographic processes in this faunal break is provided below. 

More thorough summaries for the southern and central project 

regions exist within Smith (1983), Miller and Lee (1995), Banks et al. 

(2008), and Entrix, Inc. (2008). 

2.5.1 Currents and Upwelling Events 

The driving oceanographic characteristic of the project area is the 

northerly Florida Current and the dynamics of its warm, fast-moving 

waters derived from the Gulf of Mexico and the Caribbean through the 

Yucatan Channel. In southeast Florida, coastal circulation is strongly 

related to the dynamics of the Florida Current and wave oscillations in 

the relatively narrow trough between Florida and the Bahamas 

(Soloviev et al., 2007). In the vicinity of the Jupiter and St. Lucie 

Inlets, the continental shelf platform abruptly extends wide to the east, 

deflecting the Florida Current much further offshore than in southeast 

mainland Florida (Figure 2.6), influencing the assemblages that 

utilize shallow hardbottom in manners examined in Sections 2 and 3. 

Figure 2.6. Florida Current (shown in 

orange) with offshore 

deflection near the 

southern boundary of 

Martin County, Florida 

(Image from: National 

Oceanic and Atmospheric 

Administration). 

St. Lucie Inlet 
Section 1 2-11 

Project Objectives and Nearshore Hardbottom 

within the East Florida Seascape 

Upwelling occurs in the region from wind, eddies, or current meanders that displace surface waters and 

drive replacement by cooler and usually nutrient-rich water from deeper depths (Atkinson and Targett, 

1983; Smith, 1983). The location of the western frontal boundary of the Florida Current varies 

considerably both north and south of the inflection of the current. Small frontal eddies and meanders 

propagate along the western edge of the Florida Current, typically at 1- to 2-week intervals. These 

features can transport significant nutrient and plankton patches into shallower water and contribute to 

small-scale upwelling of nutrients along midshelf and shelf break areas of this South Atlantic Bight 

subregion. Seasonal upwelling events that persist for 1 to 3 weeks typically occur during the late summer 

along some areas of east Florida (Smith, 1983). 

A high number of upwelling events occurred off north Hutchinson Island in the summer of 2003. Six 

events occurred with low temperatures ranging from 16°C to 21°C. Upwelling periods in early and late 

summer averaged 12 and 5 days, respectively (McCarthy, 2005). Analyses suggest that the longer 

upwelling events in early summer eventually resulted in similar surface and bottom temperatures. Shorter 

upwelling events during late summer appear to have resulted in stronger thermoclines because of less 

time for mixing between surface and bottom waters. Analysis of local meteorological data suggests that 

wind stress was not favorable to upwelling during most of the summer yet may have played a 

supplementary role in producing some of the upwelling events (McCarthy, 2005). In contrast to transient 

upwelling events, there is at least one area on the northern boundary of the study region where upwelling 

of nutrient-rich deep water is more permanent, an area of diverging isobaths just to the north of Cape 

Canaveral (SAFMC, in prep.). 

In Broward County, in southern portion of the project area, the Florida Current typically flows toward the 

north or northeast and the current from the surface to the bottom flows relatively consistently in the same 

direction. However, countercurrents have been recorded where the near-bottom currents flowed in a 

southerly direction (Soloviev et al., 2007). Strongest currents usually occurred near the surface, with a 

mean speed of approximately 3.4 ft/sec and a peak speed of greater than 6.8 ft/sec (Entrix, Inc., 2008). 

When the western edge of the current was closest to shore, the currents were strongest. When the 

location shifted to the east, currents sometimes weakened and for brief periods reversed and flowed 

southward. These oscillations were on the order of 14 days; however, 1- to 2-day periods were also 

recorded. 

The complexity of the system constrains full understanding of the effects of current behavior, winds, and 

upwelling on primary production and cross-shelf fluxes of nutrients and plankton. Biophysical simulations 

at mesoscale-resolution compared well with in-situ observations of shelfbreak upwelling on daily to 

seasonal timescales in east Florida (Fiechter and Mooers, 2007). Nitrate inputs associated with upwelling 

were mainly due to bottom Ekman transport as the Florida Current jet interacts with the shelf topography 

and summer winds that favor coastal upwelling. Upwelling-induced additions of nutrients and 

phytoplankton blooms can possibly increase decapod larval survival (Pitts, 1999). Brooke (2002) 

suggests that deep Oculina colonies could provide larvae to shallow-water colonies during upwelling 

events. Hare and Cowen (1996) examine a variety of physical transport mechanisms associated with 

ichthyoplankton transport and survival to the north of the study area.

2.5.2 Tides, Wind Events, Wave Activity, and Hurricanes 

The semidiurnal tides of the South Atlantic Bight vary in manners that can correspond to shelf width. The 

maximum coastal tides of 2.2 m occur at Savannah, Georgia, where the shelf is widest, and decrease to 

1.1 m at Cape Canaveral. NHB is exposed at low tides in a variety of locations, some of which are 

well-known local geologic features with intertidal habitat structure, e.g., Bathtub Reef in Martin County 

(Figure 2.7). Winds, in general, are from the northeast in fall-winter and from the southwest or southeast 

in spring-summer, but they can switch directions and form a new pattern for five or more days with little 

predictability, and with rapid increases in nearshore coastal energetics via wind and wave activity. Waves 

in the area are often wind-derived, typically from the north and northeast. Approximately half of the 

waves approach from the northeast; the largest waves approach from the north. Below northern Palm 

Beach County, wave development from the east and southeast is impeded by the Little and Great 

Bahama Banks. Banks et al. (2008) provide a recent summary of climate and wind seasonality in the 

southern and central project region. 
Section 1 2-12 

Project Objectives and Nearshore Hardbottom 

within the East Florida Seascape 

Figure 2.7. Southward view of Bathtub Reef, Stuart, Martin County, Florida. Worm rock 

(Phragmatopoma lapidosa) mounds are visible on the lower left. An aerial view of this 

intertidal reef system is shown in Figure 2.1. Photo by D. McCarthy.

During large wave events, often associated with nor’easters during fall and winter atmospheric cold 

fronts, substantial erosional events can occur. In the background, relatively typical summer-winter 

seasonal beach profiles are seen. Latitudinal differences in the impacts of these events are seen 

between Canaveral and Miami. Primary differences include 1) the very wide to very narrow north to south 

shelf width, with major shelf width change in the St. Lucie Inlet area, and 2) the positioning of the northern 

Bahama Banks off of southeast Florida, particularly from north Palm Beach County to the south. This 

wave shadow influences a variety of coastal process and sedimentary phenomena in southeast Florida. 

Hurricanes and tropical storms can produce intense wind-generated wave action during late summer and 

early fall, resulting in significant changes to the circulation in a given area. Under normal conditions, the 

Florida Current would be the dominant transport mechanism, with wave and tidal action having less 

influence. During a storm event, however, water circulation and transport are dominated by waves and 

tidal surge. During a hurricane with peak winds of at least 86 mph, the near-bottom currents on the 

inner- and mid-shelf approximately triple compared to non-storm conditions, increasing from 

approximately 0.5 to 1.5 ft/sec. During 2004 and 2005, the number of hurricane and tropical storm paths 

that impacted mainland east Florida (Figure 2.8) was higher than had been observed for many years. 

This hurricane activity may have contributed to higher post-hurricane NHB exposure in some areas 

(for example, at Singer Island, Palm Beach County, Figure 2.3). 
Section 1 2-13 

Project Objectives and Nearshore Hardbottom 

within the East Florida Seascape 

Figure 2.8. Paths of hurricanes and tropical storms impacting east Florida during A) 2005 and B) 2004. 

B) 

A) 
Section 2 

Organismal Assemblages of East Florida Shallow Hardbottom 

Section 2 

Organismal Assemblages of East Florida Shallow Hardbottom 
Section 2 3-1 

Organismal Assemblages of East Florida Shallow Hardbottom 

Chapter 3

Macroalgae and Cyanobacteria

3.1 INTRODUCTION 

NHBs along the east Florida coast provide substrate for the recruitment and growth of a number of 

species of macroalgae and cyanobacteria that serve a variety of functions. Macroalgae and 

cyanobacteria are primary producers in the marine environment and can form the foundation of a highly 

complex, diverse, and dynamic trophic food web (Duarte, 2000; Paerl, 2000; Kang et al., 2008). 

The abundance and diversity of macroalgae vary substantially within the NHB region with changes in 

latitude from Cape Canaveral to Key Biscayne. Of the varying factors along the east Florida coast that 

influence macroalgal species distribution in NHB, one of the most important likely is proximity to the 

relatively warm, oligotrophic waters of the Florida Current. Other important factors include 

species-specific life history characteristics, small- and large-scale hydrodynamics, substrate type 

(e.g., worm reefs, limestone, coral reefs) and topography, water clarity, sand scouring and burial, wave 

energy, and fish and invertebrate grazing pressure (Lobban and Harrison, 1994). 

This chapter addresses the distribution, function, and ecology of macroalgae (reds, greens, and browns) 

and cyanobacteria (blue-greens) found on NHB. Few peer-reviewed studies are available on east Florida 

coast macroalgal assemblages. Data for this review have largely been extracted from pre- or post-impact 

monitoring reports, personal communications, and personal observations made during research activities 

conducted within the study region (Appendix A, Table A.1). Macroalgae found in juvenile green turtle 

foraging studies are also listed, since these animals show high site-specificity and their foraging items are 

indicative of local macroalgal flora. Over 275 different species of macroalgae and cyanobacteria from 

108 genera within 57 families were documented from all of the studies reviewed (Appendix A, 

Table A.2). Genera and number of species within genera are also documented by county (Appendix A, 

Table A.3) in an attempt to elicit some latitudinal perspective for NHB macroalgal distribution patterns, if 

present. The algal divisions are ultimately organized into functional-form groups as proposed by Littler et 

al. (1983), which represent useful groupings of macroalgae based on morphological-metabolic-ecological 

interactions, and has been widely experimented and applied in the marine environment (Littler and Littler, 

1980; Steneck and Watling, 1982; Littler et al., 1983). 

3.1.1 Functional Form Groups 

Functional form grouping as proposed by Littler et al. (1983) assigns species of macroalgae to a 

particular group based on internal and external characteristics and is applied in this document to describe 

observed distributional patterns of macroalgae on NHB (Table 3.1). Cyanobacteria are treated 

separately in this document. The functional form algal groups are primarily based on classifying algae by 

morphological and functional adaptations, and are divided into six main groups: 1) sheet (foliose), 

2) filamentous, 3) coarsely-branched, 4) thick-leathery, 5) jointed-calcareous, and 6) crustose. The ranks 

(or groupings) are based on characteristics of evolutionary strategies that are often similar across 

divisions. For example, the thick-leathery group contains species from red, green, and brown divisions 

that are morphologically similar and share metabolic and ecological properties that can be used to 

examine their patterns or functions within a community (Littler et al., 1983). The sheet group bauplan are 

characteristically one- to two-layered (mono- or distromatic) or thin-tubed, one to several cells thick, like 

species of Ulva and Dictyota. The filamentous group contains species such as Ceramium, Spyridia, and 

Chaetomorpha, which are lightly branched, multiseriate, and lightly corticated. Macroalgal species such 

as Acanthophora, Laurencia, Liagora, Udotea, and Penicillus are placed within the coarsely-branched 

algal group, characterized as fleshy-wiry textured species with upright coarsely branched structures. 

Examples from the thick-leathery group are species of Gracilaria, Sargassum, Udotea, and Padina, 

normally several cells in thickness, toughly branched, and heavily corticated. Species in the 

jointed-calcareous group are typically articulated, calcified, and stony (e.g., Amphiroa, Jania, and 

Halimeda). Genera in the crustose group are normally characterized as encrusting, prostrate, and stony 

(e.g., Hydrolithum and Porolithon). 
Section 2 3-2 

Organismal Assemblages of East Florida Shallow Hardbottom 

Table 3.1. Macroalgal functional form group descriptions as proposed in Littler et al. (1983). 

Functional Form 

Group External Morphology Internal Anatomy Texture Genera Examined 

Sheet Thin, tubular, and 

sheet-like (foliose) 

Uncorticated, one to 

several cells thick Soft Ulva, Enteromorpha (Ulva), Dictyota

Filamentous Delicately branched 

(filamentous) 

Uniseriate, multiseriate, 

or lightly corticated Soft Centroceras, Caulerpa, Spyridia, 

Chaetomorpha, Ceramium 

Coarsely-branched Coarsely branched, 

upright Corticated Fleshy-wiry 

Acanthophora, Laurencia, 

Rhipocephalus, Liagora, Udotea, 

Caulerpa, Penicillus, Neomeris 

Thick-leathery Thick blades and 

branches 

Differentiated, heavily 

corticated, thick-walled Leather, rubbery 

Sargassum, Lobophora, Gracilaria, 

Turbinaria, Padina, Stypopodium, 

Udotea 

Jointed-calcareous Articulated, calcareous, 

upright 

Calcified genicula, 

flexible intergenicula with 

parallel cell rows 

Stony Halimeda, Galaxaura, Jania, 

Amphiroa 

Crustose Prostrate, encrusting Calcified or uncalcified 

parallel rows of cells Stony or tough Porolithon, Hydrolithon, 

Neogoniolithon, Peyssonnelia 

General differences in functional form group responses to environmental conditions (e.g., photosynthesis, 

sedimentation, grazing, etc.) are presented in Table 3.2. Data for NHB macroalgae were extracted from 

the sources listed in Appendix A, Table A.1. Macroalgae are expressed as presence/absence data 

because different sampling methods were utilized among the studies. 

Table 3.2. Effects of environmental conditions on the various algal functional form groups based on 

Littler et al. (1983). 

Functional Form 

Group Light Sedimentation Water Motion Nutrients Herbivory 

Sheet (foliose) 

High net 

photosynthesis; 

Low self-shading 

Continuous spore 

release aids in 

success 

Destructive; Some 

turf-forming species High uptake 

Low resistance 

(exceptions [e.g., 

Dictyota]); Delicate 

thalli but rapid growth 

Filamentous High net 

photosynthesis 

Commonly turf; 

Vegetation 

fragmentation aids 

in success 

Many turf-forming 

species High uptake 

Low resistance 

(exceptions [e.g., 

Caulerpa]); Delicate 

thalli but rapid and 

sometimes conspicuous 

growth 

Coarsely-branched Moderate net 

photosynthesis Commonly turf 

Turf forming species; 

Reduced 

photosynthesis near 

holdfast 

N/A 

Low to moderate 

resistance; Some 

secondary metabolic 

chemical deterrents 

Thick-leathery Moderate net 

photosynthesis 

Robust hold-fasts 

aid persistence 

Strengthens 

holdfast/reduces 

height; 

Some turf-forming 

species 

N/A High resistance; 

Morphological deterrent

Jointed-calcareous Low net 

photosynthesis Commonly turf N/A Low nutrient uptake rates 

High resistance; 

Morphological deterrent

Crustose Low net 

photosynthesis 

Tolerant of short 

burial; Anoxic 

conditions created 

under burial 

N/A Low nutrient 

uptake rates 

High resistance; 

Morphological deterrent

N/A = not available. 
Section 2 3-3 

Organismal Assemblages of East Florida Shallow Hardbottom 

3.1.2 Latitudinal Gradients 

There are several dramatic and some relatively subtle differences in macroalgal distribution along east 

Florida NHB that are most likely correlated to two major geographical features: 1) the diverging path of 

the Florida Current (or Gulf Stream), and 2) gradients in seawater temperatures from warm temperate to 

subtropical latitudes. The average distance of the Gulf Stream from the nearshore waters of south 

Florida is approximately 24 km, while the average distance in the northern NHB is approximately 48 km 

away at Port Canaveral (Figure 3.1). Between Cape Canaveral and Palm Beach, Searles (1984) 

documented the southernmost limit of several warm temperate species (e.g., Hypnea volubilis, 

Petroglossum undulatum, Gloioderma blomquistii) and the northern limits of a number of subtropical 

species. Macroalgal biomass appears to be relatively higher in northern east Florida coast NHB counties 

than in the southern NHB counties. While the specific reasons for this have not been experimentally 

tested, the trend may be due to latitudinal differences in nutrients (possibly a function of the oligotrophic 

waters of the Florida Current), wave dynamics, and abundance and diversity of grazing fishes and 

invertebrates. The later explanation may be further supported by the relatively higher percentage of 

physically- and chemically-defended macroalgae (i.e., thick-leathery, jointed-calcareous, and crustose 

forms) seen towards southeast Florida (see Chapter 7) and, possibly, a relatively higher abundance and 

diversity of herbivorous fish species seen towards more southern latitudes, as suggested by Floeter et al. 

(2004). Presence/absence data from NHB studies support some of these general observations 

(e.g., increase in defended species, especially crustose and jointed-calcareous); however, these data 

have not been adequately tested. 

Figure 3.1. Path of the Florida Current (Gulf Stream). Note the increased distance of the Gulf Stream 

west wall along the east Florida coast from south to north (Image from: Naval 

Oceanographic Office website [unclassified document]). 
Section 2 3-4 

Organismal Assemblages of East Florida Shallow Hardbottom 

3.1.3 Depth Gradients 

The overall percent compositions of red (Rhodophyta), green (Chlorophyta), and brown (Phaeophyta) 

macroalgal species documented in NHB studies (Figure 3.2) appear to reflect the overall taxonomic 

abundance of species of the different divisions. For example, red algae represent the largest taxonomic 

group of macroalgae, and red algae composed the highest percent composition of total NHB macroalgae. 

Saffo (1987) and others have more recently questioned Engelmann’s theory (1883; see Saffo, 1987) on 

the depth distributions among green, brown, and red macroalgae. The more-than-century-old hypothesis 

that predicted chromatic adaptation phylogeny was responsible for the vertical distribution of macroalgae 

has been tested with more advanced technology. Biologists now recognize that factors other than 

photosynthetic pigments may affect the distribution of macroalgae, and that seaweeds photoacclimate to 

light quality as well as light intensity (Saffo, 1987; Dawes, 1998). 

5.33%

24.85%

15.68%

54.14%

Cyanobacteria

Chlorophyta

Phaeophyta

Rhodophyta

Figure 3.2. Percent composition of macroalgae and cyanobacteria divisions documented from studies 

conducted on nearshore hardbottom (see Appendix A, Table A.1). 

Macroalgae from the functional form groups of filamentous and thick-leathery represented about 50% of 

the macroalgae documented from studies conducted on east Florida NHB (Figure 3.3). Functional form 

group representation, however, varied among counties (Figure 3.4). Combinations of several different 

local microhabitat features (e.g., wave dynamics, substrate type, turbidity levels) are most likely 

responsible for macroalgal zonation patterns of diversity and abundance, as well as changes in form 

(turfing) on NHB. Turf-forming algae, defined here as macroalgae that form colonial assemblages instead 

of spatially segregated individuals, normally form in stressful environments, such as areas that 

experience periodic or frequent sedimentation, exposure, and/or high grazing pressure (Airoldi, 2001). 

Conditions that induce algal “turfing” are common in low intertidal and shallow subtidal areas. The 

hardbottoms in these areas are usually well-vegetated with fast-growing and/or turf-forming macroalgae, 

especially on high-energy beaches. Some patterns elicited among the functional form groups are 

1) resilience among many species in the sheet and filamentous groups, and 2) resistant species in the 

thick-leathery group. Distributional patterns indicate the morphology and productivity strategies of 

articulated coralline and crustose functional form species enable them to either survive in stressful 

environments or select against settlement in these habitats. 
Section 2 3-5 

Organismal Assemblages of East Florida Shallow Hardbottom 

9%

30%

24%

21%

9%

1% 6%

Sheet

Filamentous

Coarsely-Branched

Thick Leathery

Jointed Calcareous

Crustose

Unid. & Cyanobacteria

Figure 3.3. Percent composition of macroalgae and cyanobacteria functional form groups documented 

from studies conducted on nearshore hardbottom (see Appendix A, Table A.1). 

0%

5%

10%

15%

20%

25%

30%

35%

40%

45%

Brevard Indian

River

St. Lucie Martin Palm

Beach

Broward

Sheet 

Filamentous

Coarsely Branched 

Thick Leathery 

Jointed Calcareous

Crustose 

Unid

Figure 3.4. Percent composition of macroalgal functional form groups (by county) documented from 

studies conducted on nearshore hardbottom (see Appendix A, Table A.1). 

3.1.4 Algal Reproduction 

3.1.4.1 Sexual Reproduction 

Most macroalgae are capable of sexual and asexual reproduction; however, the latter remains less costly. 

Sexual reproduction promotes variation at the cost of sporophyte and gametophyte production. Thus far, 

research on the reproductive strategies of certain macroalgae indicates there are several variations within 

and between species, and generalizations should be cautioned (Lobban and Harrison, 1994). However, 

for descriptive purposes, a brief definition of the basic reproductive strategies is made here to apply their 

ecological significance to NHB macroalgae. The three basic life history strategies (from Moore et al., 

1998) are as follows: 
Section 2 3-6 

Organismal Assemblages of East Florida Shallow Hardbottom 

• sporic meiosis – diploid parents form spores by sporic meiosis; spores produce haploid parents, 

which produce gametes (i.e., Ulva lactuca, U. flexuosa [formerly Enteromorpha], Polysiphonia spp., 

Laurencia spp.). 

• gametic meiosis – where diploid parents directly produce gametes by gametic meiosis, bypassing the 

haploid parent phase (i.e., Sargassum spp., Turbinaria spp.). 

• zygotic meiosis – where haploid parents produce gametes, and gametes fuse into a zygote, which is 

the only diploid cell in this life cycle (i.e., Cladophora spp., Ulva spp.). 

Heteromorphic or isomorphic haplodiplontic sporic meiosis is a common life history strategy of many NHB 

macroalgae. Some advantages of this life history (two free-living phases) are that there are different 

photosynthetic characteristics between gametophytes and sporophytes. There are shade-adapted young 

sporophytes and gametophytes and sun-adapted (canopy) mature sporophytes (Littler and Littler, 1980 in 

Dawes, 1998). Other advantages of this life history may include grazer-resistant diploid crustose phases 

under heavy herbivory levels that alternate with bladed haploid phases when under reduced herbivory 

pressure (Dawes, 1998). 

The dispersal distances of algal propagules vary considerably with species (i.e., 3 m versus 500 m; see 

Reed et al., 1988) due to species life history in combination with episodic events (i.e., storm events). 

However, most macroalgae seeds generally settle within a relatively short range (~1 m) from their source 

(Deysher and Norton, 1982; Lobban and Harrison, 1994). Research to predict rates and distances of 

dispersal in benthic marine communities has been a recent focus in the marine ecology field. Carr et al. 

(2003) contends that marine systems are more “open” and dispersal is relatively wider than in terrestrial 

systems. However, Cowen et al. (2000) reviewed several studies that indicate small-scale circulations in 

marine systems tend to minimize long-distance dispersal, creating higher-than-expected retention of 

invertebrate larvae (Cowen et al., 2000) and macroalgal propagules (Santelices, 1990). Bobadilla and 

Santelices (2005) constructed a model of the predicted spatial distribution of macroalgal dispersal with 

expected high rates of settlement near the parent plant and progressively less settlement with increasing 

distance from the propagule source. However, models run with in-field propagule data did not conform to 

the expected results, with actual propagule dispersal rates being pulsed and highly variable. Source 

curves that were generated from their data indicated dispersal distance was greater from the source than 

predicted for many rocky intertidal species. Ultimately, results from Cowen et al. (2000) and Bobadilla 

and Santelices (2005) support the idea that more accurate models need to be derived that include 

1) localized currents and eddies and 2) pulsing events in the capacity of propagules to be imported and 

exported. Additionally, another important factor to consider is that dispersal distances and survival rates 

can be enhanced by consumers, where spores successfully pass through the gut of gastropods or fishes 

and benefit from the sinking, sticking, and nutrient qualities of the fecal pellet (Santelices and Paya, 

1989). 

3.1.4.2 Asexual Reproduction 

Several different strategies exist for clonal (asexual) reproduction: 1) stoloniferous growth (e.g., Caulerpa

spp. and Penicillus spp.), where the plant consists of a prostrate stolon and upright growth, 2) common 

holdfast growth (e.g., Corallina officinalis and Ascophyllum spp.), where a basal holdfast gives way to 

multiple upright thalli, 3) crusts (e.g., Mesophyllum spp. and Ralfsia spp.), which grow out horizontally, 

4) the bending and reattaching of plants to the substrate (e.g., Laurencia spp.), where the apical cell 

attaches to the surface and generates a new thallus, and 5) fragmentation of parts (e.g., Halimeda spp., 

Dictyota spp., Gracilaria spp.) (Collado-Vides, 2002). 

3.1.4.3 Life History Strategy (Perennial vs. Annual Algae) 

There are a number of life history strategies of macroalgae, but most can be categorized as either 

perennial (live for multiple years) or annual (live for 1 year). Perennial algae often represent the final 

stages of succession and are capable, in some cases, of living for up to 20 years (Lüning, 1990). 

Perennials exist in, and are dependent on, stable environmental conditions (or at least predictable 

fluctuating conditions). They are also resilient to low light and nutrient marine environments. Perennial 
Section 2 3-7 

Organismal Assemblages of East Florida Shallow Hardbottom 

algae can be categorized as k-strategist species that optimally utilize environmental resources. Their 

annually fixed organic carbon stores are used to increase thallus growth (e.g., large canopy species) in 

lieu of allocating resources toward reproductive structures. In contrast, the biomass of annual species 

(r strategy) is highly variable since they allocate much of their resources to generate reproductive cells on 

the thallus and reproduce quickly (Lüning, 1990). Macroalgal species associated with the NHB in Florida 

appear to have greater numbers of annual species in intertidal and shallow subtidal waters than in deeper 

(>31 m) waters off the Florida coast, where most of the documented macroalgal species are considered 

perennial (Hanisak and Blair, 1988). 

3.1.5 Functional Form Group Descriptions 

3.1.5.1 Sheet Group 

Littler et al. (1983) described this group as having a soft, foliose, and uncorticated external morphology 

with a general cell thickness that ranges from one to several cells. Representative genera from this group 

are Ulva and Dictyota (Figure 3.5). Predominantly fast-growing and found in shallow waters, species in 

this group typically have high net photosynthesis rates, are short-lived, and are nitrogen-limited (Littler 

and Arnold, 1982). 

Figure 3.5. Representative species of the sheet functional form group growing on nearshore 

hardbottom: Ulva lactuca (left) and Dictyota sp. (right). Photos by K. Holloway-Adkins. 

Thirty-one different species based from studies conducted on east Florida NHB were from the sheet 

functional form group: 8 green (Ulva spp.), 22 brown (Dictyopteris delicatula and Dictyota spp.), and 1 red 

macroalga (Nitophyllum punctatum). Macroalgae within this functional group were highly represented in 

Martin and Brevard Counties (Figure 3.6). The number of sheet group species among NHB studies 

ranged from 0% (Salmon et al., 2004) in Broward County to 20% (Cummings, 1990) in Palm Beach 

County (Appendix A, Table A.4). The number of sheet group species was highest in foraging sample 

analyses of juvenile green turtles in Palm Beach County (27.3%) (Makowski, 2004) (Appendix A, 

Table A.4). 

The sheet group is abundant throughout east Florida NHB and can be found in both intertidal and subtidal 

waters. Species of Ulva and Dictyota were the most frequently represented species from this group and 

were documented in all counties within the NHB area. Species like Ulva that are thin and transparent are 

less efficient in absorbing submarine illumination than thick, multilayered plants like Codium, even though 

they share the same pigments (Dawes, 1998). However, light can be effectively trapped at depth through 

morphological adaptations and changes in pigment concentration. 
Section 2 3-8 

Organismal Assemblages of East Florida Shallow Hardbottom 

Percentage of Sheet Group by County

13.16%

7.45%

8.33%

14.29%

7.79%

12.71%

0.00%

2.00%

4.00%

6.00%

8.00%

10.00%

12.00%

14.00%

16.00%

BREVARD INDIAN

RIVER

ST. LUCIE MARTIN PALM BEACH BROWARD

Figure 3.6. Percentage of species in the sheet functional form group documented in studies conducted 

on nearshore hardbottom in counties from Brevard to Broward, Florida (see Appendix A, 

Table A.4). 

Most of the green and brown algae within the sheet group display haplodiplontic sporic meiosis life 

history. Littler et al. (1983) found net productivity was high (second highest of groups) for macroalgae 

characterized within the sheet group. Species are likely to be opportunistic, with limited life history 

strategy flexibility (Littler and Littler, 1980); however, because most species from this group have long 

continuous spore release events (Eriksson and Johansson, 2005), they are highly productive. Species 

from this group are commonly early successional species, short-lived, and very resilient to environment 

disturbances. 

Many of the sheet group species have bauplans with high surface-to-volume ratios, which promotes a 

higher uptake of nutrients and accelerates photosynthesis. The ubiquitous Ulva was described by 

Guimaraens and Coutinho (2000) as light-limited during winter and nutrient-limited during summer. 

However, upwelling events that occur in summer, coupled with the effects of high irradiance and excess 

nutrients, significantly stimulated algal growth of this species. The sheet group tends to be opportunistic 

(r strategy), settle, and grow quickly, and be eurytopic (tolerant of a wide range of environmental 

conditions or habitats); however, this group is more prone to desiccation and grazer damage. Most of 

these forms are comparatively more susceptible to damage from sedimentary particle abrasion and the 

shearing action of high-energy waves. Species from this group (i.e., Ulva and Dictyota) were frequently 

found in foraging analyses of juvenile green turtles and herbivorous fishes (see Chapter 7). However, 

experimental studies indicate that compounds inherent to Dictyota may deter many grazing invertebrates 

and fishes (Cronin et al., 1997). 

3.1.5.2 Filamentous Group 

The morphology of this group has been described as delicately branched and filamentous with internal 

anatomy characterized as uniseriate, multiseriate, or lightly corticated (Littler et al., 1983). Typical genera 

include Centroceras, Chaetomorpha, and Ceramium. This group represented the largest functional form 

group (31.0%) found on east Florida NHB and was documented from studies in every county examined. 

Ninety-seven macroalgal species were identified as having filamentous functional form. This group 

included 24 green, 9 brown, and 64 red algae. Most of the same species were abundant and common 

along the entire east Florida NHB; however, they were most frequently documented in Indian River and 

Brevard Counties (Figure 3.7). The number of filamentous group species for NHB studies ranged from 

0% (Continental Shelf Associates, Inc., 2005b) in Palm Beach County to 35.8% (Juett et al., 1976) in 

Indian River County (Appendix A, Table A.5). The highest composition of filamentous species found in 

foraging analysis of juvenile green turtles was 36.3% in Palm Beach County (Makowski, 2004) 

(Appendix A, Table A.5). 
Section 2 3-9 

Organismal Assemblages of East Florida Shallow Hardbottom 

Percentage of Filamentous Group by County

31.58% 32.92%

27.78%

30.61%

18.18% 19.49%

0.00%

5.00%

10.00%

15.00%

20.00%

25.00%

30.00%

35.00%

BREVARD INDIAN

RIVER

ST. LUCIE MARTIN PALM

BEACH

BROWARD

Figure 3.7. Percentage of species in the filamentous functional form group documented in studies 

conducted on nearshore hardbottom in counties from Brevard to Broward, Florida 

(see Appendix A, Table A.5). 

Macroalgae from the filamentous form group were documented among most of the east Florida NHB 

studies. Bryopsis, Caulerpa, and Ceramium were reported in studies within all of the counties, and 

Cladophora was counted in all but Broward County (Appendix A, Table A.3). These species are present 

and abundant in both intertidal and subtidal waters (K. Holloway-Adkins, East Coast Biologists, Inc., 

[ECB], personal observation) and exhibit few inhibitions to growth or distribution. Many species within this 

group are frequently found in the diet of herbivorous invertebrates, fishes, and juvenile green turtles. 

Net productivity was experimentally highest for macroalgae within the filamentous functional form group 

(Littler et al., 1983). Most macroalgae in this group exhibit high tolerance for changes in sedimentation 

and will commonly turf to persist in stressful environmental conditions. Dense turfs of filamentous algae 

are common in lower shallow intertidal waters, and this morphological adaptation helps these species 

persist and grow under intense grazing pressure and/or high-energy beach conditions (Airoldi, 2001). 

A few species can grow epiphytically on other macroalgae; however, most form a somewhat distinct 

holdfast or stolon and require hard substratum for attachment and growth. Most of the species within this 

group lack defense metabolites and are frequently consumed by many herbivorous fishes, invertebrates, 

and juvenile green turtles (see Chapter 7). 

Green and brown filamentous group macroalgae exhibit haplodiplontic sporic meiosis life history 

strategies. The majority of species in this group elicit r-strategy characteristics and are highly resilient but 

short-lived species. Reproductive strategies of red algae within this group are known to have long 

continuous spore releases events (Eriksson and Johansson, 2005). Additionally, species from this group 

are commonly early successional species, highly ephemeral, and characteristically resilient. 

3.1.5.3 Coarsely-branched Group 

The morphology of this group has been described by Littler et al. (1983) as coarsely-branched, upright, 

and corticated. Typical genera include species of Acanthophora, Laurencia, Liagora, Udotea, Caulerpa, 

and Penicillus; Laurencia and Caulerpa are pictured in Figure 3.8. 
Section 2 3-10 

Organismal Assemblages of East Florida Shallow Hardbottom 

Figure 3.8. Representative species of the coarsely-branched functional form group growing on 

nearshore hardbottom: Laurencia sp. (left image, top left and right) and Caulerpa sp. (cover 

in right image). Photo by K. Holloway-Adkins.

There were 83 macroalgal species within the coarsely-branched functional form group documented from 

east Florida NHB studies. The group itself is dominated by species of red and green macroalgae (see 

Littler et al., 1983). Fifty-seven species of red, 26 species of green, and no species of brown macroalgae 

were documented in NHB studies for this functional form group. 

The percent of macroalgae from this group was highest in Brevard County (39%; Figure 3.9). The 

number of coarsely-branched species per NHB study ranged from 16.7% composition in Broward County 

(Salmon et al., 2004) to 66.7% in Palm Beach County (Continental Shelf Associates, Inc., 2005b) 

(Appendix A, Table A.6). The percent composition of macroalgae from the coarsely-branched functional 

group was highest in the foraging samples of juvenile green turtles in Broward County (Wershoven and 

Wershoven, 1992a) at 40.0% (Appendix A, Table A.6). 

Percentage of Coarsely-Branched Group by County

39.47%

22.98%

29.86%

22.45%

32.47%

26.27%

0.00%

5.00%

10.00%

15.00%

20.00%

25.00%

30.00%

35.00%

40.00%

45.00%

BREVARD INDIAN

RIVER

ST. LUCIE MARTIN PALM

BEACH

BROWARD

Figure 3.9. Percentage of species in the coarsely-branched functional form group documented in 

studies conducted on nearshore hardbottom in counties from Brevard to Broward, Florida 

(see Appendix A, Table A.6). 

This group contains species that extend to the edge of the range and therefore exhibit traits from the two 

adjacent groups, i.e., on one end some genera like Udotea have low net productivity and tough external 

morphology, but others share other characteristics with the coarsely-branched group (Littler et al., 1983). 
Section 2 3-11 

Organismal Assemblages of East Florida Shallow Hardbottom 

Acanthophora spicifera exhibits seasonal patterns of growth and mortality on east Florida NHB 

(K. Holloway-Adkins, ECB, pers. observ.); however, in tropical Panama it was considered perennial (Kilar 

and MacLachlan, 1989). Some species within this group have dormant or reduced stages where the 

plant dies back to the holdfast and regenerates under more favorable (i.e., photoperiod, temperature, 

salinity level, etc.) conditions (Brawley and Johnson, 1992). 

3.1.5.4 Thick-leathery Group 

The morphology of this group is described as having thick blades and branches, with internal anatomy 

being differentiated, heavily corticated, and thick-walled (Littler et al., 1983). Species include Sargassum, 

Lobophora, Gracilaria, Turbinaria, Padina, Stypopodium, and Udotea. 

A total of 71 different species was documented from east Florida NHB studies (38 red, 21 brown, and 

12 green macroalgae). Species within this group were widely distributed across the NHB in every county 

and were from all three macroalgal divisions. Gracilaria and Padina occur frequently throughout east 

Florida NHB, while Sargassum, Lobophora, Turbinaria, and Udotea are more frequently found in the 

central and southern east Florida NHB. Representative species from the thick-leathery functional form 

group are shown in Figure 3.10. 

Figure 3.10. Several representative species of the thick-leathery functional form group, including 

Agardhiella subulata and Gracilaria sp., growing on nearshore hardbottom. Photo by 

K. Holloway-Adkins.

Distribution – Latitude and Water Depth 

The percent of macroalgae from this group was highly represented in Broward and Martin Counties 

(Figure 3.11). Percent composition found in studies conducted along east Florida NHB showed a low 

percentage (8%) in Palm Beach County (Continental Shelf Associates, Inc., 2005b) and a high 

percentage (50%) in Broward County (Salmon et al., 2004) (Appendix A, Table A.7). Species from the 

thick-leathery group were highest (30.0%) in foraging samples that were analyzed from juvenile green 

turtles in Broward County (Wershoven and Wershoven, 1992b) (Appendix A, Table A.7). 
Section 2 3-12 

Organismal Assemblages of East Florida Shallow Hardbottom 

Percentage of Thick-Leathery Group by County

13.16%

19.88%

23.61% 24.49%

16.88%

24.58%

0.00%

5.00%

10.00%

15.00%

20.00%

25.00%

30.00%

BREVARD INDIAN

RIVER

ST. LUCIE MARTIN PALM BEACH BROWARD

Figure 3.11. Percentage of species in the thick-leathery functional form group documented in studies 

conducted on nearshore hardbottom in counties from Brevard to Broward, Florida 

(see Appendix A, Table A.7). 

Robust holdfast structures may promote persistence under short-term burial events. This group’s 

reproductive strategy appears to be not as tolerant of sedimentation (Eriksson and Johansson, 2005); 

however, their association with turf (understory) species may promote their persistence in stressful 

environments (Johnson and Brawley, 1998). The majority of species within this group produce secondary 

metabolites and persist as perennial k-strategy plants. 

3.1.5.5 Jointed-calcareous Group 

External morphology of this group is described as being articulated, calcareous, and upright (Littler et al., 

1983). Internal anatomy is calcified genicula and flexible intergenicula with parallel cell rows. Species 

include Halimeda, Galaxaura, Jania, and Amphiroa. Red coralline species (Order Corallinales) are less 

frequently encountered in the northern reach of the east Florida NHB but are represented by Amphiroa

sp. and Jania spp. throughout the entire east Florida NHB. 

There were 32 different species (19 rhodophytes and 13 chlorophytes) from the jointed-calcareous 

functional form group documented from east Florida NHB studies. Jointed-calcareous functional form 

group species were represented in every county within the NHB. 

Broward County had the highest percent of macroalgae from this group (Figure 3.12). Percent 

composition found in studies conducted along east Florida NHB were highest (21.7%) in the Vare study 

(1991) in Palm Beach County (Appendix A, Table A.8). The highest percent composition of 

jointed-calcareous species found in foraging samples of juvenile green turtles was 10%, in Broward 

County (Wershoven and Wershoven, 1992b) (Appendix A, Table A.8). 
Section 2 3-13 

Organismal Assemblages of East Florida Shallow Hardbottom 

Percentage of Jointed-Calcareous Group by County

2.63%

5.59%

6.94%

4.08%

14.29%

16.10%

0.00%

2.00%

4.00%

6.00%

8.00%

10.00%

12.00%

14.00%

16.00%

18.00%

BREVARD INDIAN

RIVER

ST. LUCIE MARTIN PALM

BEACH

BROWARD

Figure 3.12. Percentage of species in the jointed-calcareous functional form group documented in 

studies conducted on nearshore hardbottom in counties from Brevard to Broward, Florida 

(see Appendix A, Table A.8). 

Calcareous species are important contributors to sediment production (as well as species from the 

crustose group). They exhibit more k-strategy life histories, transferring energy towards structural 

development (e.g., calcareous external morphology) and therefore growing much slower than filamentous 

and sheet group species. Net productivity of this group and the crustose group are lowest among the 

functional form groups (Littler et al., 1983). The propagules of articulated coralline species are 

considered poor dispersers (Dethier et al., 2003), and macroalgal dispersal generally serves to primarily 

replenish local productivity. 

Parrotfish (Scaridae) readily consume calcareous algae (e.g., Halimeda spp., Avrainvillea spp., 

Penicillium spp.). However, it appears that other herbivorous fish species and most invertebrates avoid 

calcareous green and red algal species. 

3.1.5.6 Crustose Group 

Morphology for this group is typically epilithic, prostrate, or encrusting. The internal morphology has been 

described as calcified or uncalcified parallel rows of cells (Littler et al., 1983). From this group, only 

species of Rhodophyta were found on east Florida NHB. 

The crustose macroalgal group was represented in only two counties, so a multi-county figure similar to 

Figures 3.6 through 3.12 is not presented. Lithophyllum sp. and Peyssonnelia rubra were the only two 

species documented on east Florida NHB. Lithophyllum sp. was documented in a study in Broward 

County (USACE, 2003a), and Peyssonnelia rubra was documented by Juett et al. (1976) in Indian River 

County. 

Studies conducted showed a high percentage (18.5%) for crustose-group in Broward County (USACE, 

2003a) and a low percentage (1.8%) in Indian River County (Juett et al., 1976) (Appendix A, Table A.9). 

Foraging analysis of juvenile green turtles did not include any species in the crustose functional form 

group. 

Species within this group show low net productivity; however, they are highly resistant to mechanical 

damage by sedimentation events and grazers, are considered perennial species, and can reproduce both 
Section 2 3-14 

Organismal Assemblages of East Florida Shallow Hardbottom 

asexually and sexually. Species in this group can grow in deep intertidal and shallow subtidal areas of 

the reef (as suggested by van den Hoek et al., 1975 in Dawes, 1998). 

3.1.5.7 Cyanobacteria 

Cyanobacteria are photosynthetic, oxygen-producing bacteria that were formerly known as blue-green 

algae. They play a major role in nitrogen fixation and are primary producers in most marine 

environments. Some occur at macroscopic sizes (i.e., Lyngbya), but most are microscopic. The 

paramount importance of cyanobacteria in shaping the supporting marine food webs is sometimes 

underestimated (Paerl, 2000). In some cases, cyanobacteria have been reported to produce up to 

20% of ocean primary production (Moore et al., 1998), while others report it to account for anywhere from 

30% to over 50% of the global oceanic primary production (Paerl, 2000). Their ubiquitous presence, 

eurythermal nature, and, sometimes, dual life-history strategy (auto- and heterotrophic) have made them 

difficult to classify, as well as characterize. Most likely several hundred different species are associated 

with NHB on Florida’s east coast. 

The two basic morphological classifications for marine planktonic cyanobacteria are 1) unicellular, and 

2) filamentous. Filamentous forms are further subdivided into heterocystous and non-heterocystous 

forms (Paerl, 2000). Examples of heterocystous filamentous cyanobacteria documented from NHB 

studies were Anabaena and Nodularia. Non-heterocystous blue-green algae found included genera of 

Lyngbya and Oscillatoria. 

Except for the more epiphytic colonial, filamentous cyanobacteria, most species are microscopic in size. 

Cyanobacteria can be relatively difficult to identify in situ and are predominantly inconspicuous in most 

cases. The paucity of data on species found on NHB from reviewed studies is most likely due to 

blue-green algae being overlooked and/or undocumented. Ongoing research by the Smithsonian Marine 

Station in Fort Pierce, Florida and Nova Southeast University in Ft. Lauderdale is exploring the 

cyanobacteria interactions with other sessile organisms along the Florida coast. For example, their 

research suggests that cyanobacteria blooms appear to be more frequent along the Florida coast in 

recent years and may prohibit coral recruitment (Kuffner et al., 2006; Paul et al., 2007). Species of 

cyanobacteria were recorded for Indian River (Juett et al., 1976), Martin (Phillips, 1961), and Palm Beach 

(Vare, 1991) Counties, and from foraging samples of juvenile green turtles in Palm Beach County (Jones 

et al., 2004) (Appendix A, Table A.1). High-density Lyngbya blooms were documented on Gulf Stream 

reef (Boynton Beach OHB) from 2002 to 2004 and again in fall 2008 by Palm Beach County Reef 

Rescue. Also, Lyngbya blooms occurred on Broward County NHB and OHB from 2002 through 2004, 

where they smothered benthic species, particularly octocorals (Paul et al., 2007; Banks et al., 2008). In 

addition, NHB in Broward County south of Port Everglades was dominated by cyanobacteria during the 

summer of 2001 (Miller and Cuba, 2003; USACE, 2003a). 

Seventeen species from 11 genera and eight different families were documented from east Florida NHB 

studies (Appendix A, Table A.2). The most frequently documented species were Calothrix, Lyngbya, 

Microcoleus, and Schizothrix. Other species observed were Anabaena, Anacystis, Entophysalis, 

Hormothamnion, Nodularia, Oscillatoria, and Porphyrosiphon.

There is very little information available on how diversity of cyanobacteria changes along the NHBs of the 

east Florida coast. However, it can be fairly easily observed in both intertidal and subtidal hardbottoms, 

particularly during the late summer time period. Peaks in abundance of an unknown filamentous 

cyanobacteria species on dead intertidal worm reefs during late summer have been consistently observed 

(D. McCarthy, Jacksonville University, Jacksonville, FL, pers. observ.). Subtidally, it also appears that 

filamentous cyanobacteria (likely Lyngbya sp.) may bloom during the summer time period. Peaks in 

abundance may be enhanced during this time period when late summer upwelling events provide excess 

nutrients to these shallow waters (McCarthy, 2005). 

Cyanobacteria reproduce asexually and possibly via parasexual processes. Asexually, filamentous forms 

reproduce by trichome fragmentation, or by formation of special hormogonia (distinct reproductive 

segments of the trichomes). Hormogonia display gliding motions and gradually develop into new 
Section 2 3-15 

Organismal Assemblages of East Florida Shallow Hardbottom 

trichomes (World Health Organization, 1999). They also contain akinetes, which are modified vegetative 

cells that can remain dormant for long periods of time until environmental conditions are favorable to the 

germination of the new filaments. While there is no evidence of sexual reproduction, cyanobacteria may 

reproduce parasexually via transformation and conjugation (like bacteria). 

3.2 SUMMARY 

NHB areas off Florida’s east coast provide critical substrate for the settlement, growth, and production of 

macroalgae and filamentous cyanobacteria. Some important functions of macroalgal communities are to 

1) provide oxygen through respiration processes, 2) aid in nutrient cycling, and 3) provide food and 

shelter for numerous species of herbivorous fishes, invertebrates, and juvenile green turtles. For 

example, the chemical content and architecture of Laurencia sp. function as important cues for spiny 

lobster postlarval settlement and subsequent juvenile shelter selection. Both use the architecture of, and 

the food present on, Laurencia in selecting habitat (Herrnkind and Butler, 1986). Other macroalgal 

species such as the ubiquitous calcareous Halimeda also contribute three-dimensional structure for many 

species while alive, as well as calcium carbonate sands after they die and decompose. In particular, in 

the Florida Keys the contribution of Halimeda is widely evident where the process of deposition and 

sedimentation enrich local substrata, promoting the growth and productivity of other macroalgal and 

invertebrate species (Wiman and McKendree, 1975). 

There have been several noted invasive magroalgal species encountered along Florida’s NHB. 

Generally, Williams and Smith (2007) report that there currently are 277 species of macroalgae that have 

been introduced at the global scale due to human activities. Those with opportunistic life history 

characteristics make them more likely to become invasive species under certain conditions 

(i.e., siphonous greens like Caulerpa spp.) and consequently warrant a higher level of monitoring 

(Williams and Smith, 2007). Generally, most macroalgae are nitrogen-limited, and when land-based 

runoff creates high nutrient levels, macroalgal proliferation can increase. Under these conditions, some 

non-native as well as native species can become invasive monocultures that smother corals and other 

invertebrates, limiting availability light and creating anoxic conditions. LaPointe et al. (2005) investigated 

N sources predominantly responsible for an invasive species outbreak (Caulerpa brachypus) and the 

exponential production of a native species (Codium isthmocladum) that became harmful algal blooms in 

Palm Beach and Broward County. Through “signature” analysis of nitrogen, they determined the greatest 

source of available and assimilable nitrogen was from land-based sewerage systems. 

Macroalgae can be classified according to morphological and functional adaptations and are divided into 

six main groups: 1) sheet (foliose), 2) filamentous, 3) coarsely-branched), 4) thick-leathery, 

5) jointed-calcareous, and 6) crustose. 

While no one particular alga would completely align with the definition of keystone species, there are a 

handful of species within the macroalgal community that are both abundant on NHB and play an 

important role among the grazers. Ten genera (Caulerpa, Ceramium, Codium, Dictyota, Gelidium, 

Graciliaria, Laurencia, Padina, Ulva, Bryopsis) recurred in NHB studies for all six counties examined. 

Ceramium, Ulva, Dictyota, Hypnea, Laurencia, Gelidium, Caulerpa, and Jania were found in the foraging 

samples of herbivores, and were documented in studies in at least five of the six counties. Some general 

patterns observed for macroalgae in different NHB regions (counties) were 1) no calcareous green 

(including Halimeda) species were found in Brevard County, 2) fewer Caulerpa species were present 

north of Sebastian Inlet (Brevard County) when compared to NHB in counties south of this area, 

3) a higher number of red algal species in northern NHB counties (Indian River and Brevard), and 

4) relatively few species of brown algae in northern NHB. 

While there are not a great deal of comparable macroalgal data from along the east Florida coast, 

biomass appears to be relatively higher in the northern east Florida coast NHB counties than in the 

southern ones. While the specific reasons for this have not been experimentally tested, the trend may be 

due to latitudinal differences in nutrients (possibly a function of oligotrophic waters of the Florida Current), 

wave dynamics, and abundance and diversity of grazing fishes and invertebrates. Finally, cyanobacteria 

can be commonly encountered along the east Florida NHB, although there is not much known about how 
Section 2 3-16 

Organismal Assemblages of East Florida Shallow Hardbottom 

diversity changes with depth and latitude. Ongoing research at two major institutions on the east Florida 

coast is exploring community interactions involving cyanobacteria and sessile organisms in subtropical 

ecosystems. 

Macroalgae from the filamentous and thick-leathery functional form groups represented more than 50% of 

the macroalgae documented from studies conducted on east-Florida NHB. Ninety-seven macroalgal 

species were identified as having filamentous functional forms. This group included 24 green, 9 brown, 

and 64 red algae. Commonly encountered thick-leathery group (thick blades and branches with 

differentiated internal anatomy heavily corticated, and thick body walls) species included Sargassum, 

Lobophora, Gracilaria, Turbinaria, Padina, Stypopodium, and Udotea. In contrast, there were 31 species 

from the sheet functional form group that included 8 green (Ulva sp.), 22 brown (Dictyopteris delicatula

and Dictyota sp.), and 1 red (Nitophyllum puncatum) macroalgal species. There were 75 macroalgal 

species within the coarsely-branched group, which was dominated by red and green algae with typical 

genera of Acanthophora, Laurencia, Liagora, Udotea, Caulerpa, and Penicillus. Common species from 

the jointed-calcareous group (articulated, calcareous, and upright) include Halimeda, Galaxaura, Jania, 

and Amphiroa. Within this group, macroalgae on the hardbottom in Brevard/Indian River Counties 

showed no Halimeda north of Sebastian Inlet, relatively few Caulerpa species, and no calcareous greens 

when compared to NHB in counties south of this area. Finally, crustose group members such as 

Peyssonnelia could be encountered but are not well-documented in Florida NHB. 

Generally, there appears to be greater numbers of annual species in intertidal and shallow subtidal 

waters along the east Florida coast than in deeper (>31 m) waters off the Florida coast, where most of the 

documented macroalgal species are considered perennial. Some patterns elicited among the functional 

form groups encountered along the east Florida coast are 1) resilience among many species in the sheet 

and filamentous groups, and 2) resistant species in the thick-leathery group. In the shallower, harsher 

NHB, most macroalgae in the filamentous functional group exhibit high tolerance for changes in 

sedimentation and will commonly turf to persist in stressful environmental conditions. Dense turfs of 

filamentous algae are common in lower shallow intertidal waters, and this morphological adaptation helps 

these species persist and grow under intense grazing pressure and/or high-energy beach conditions. In 

contrast, the sheet group tends to be opportunistic (r strategy), settle, and grow quickly, be eurytopic in 

nature, and incur high levels of physical damage from grazing and sedimentation activities. 

Regardless of general life history strategy, most macroalgae along the east Florida NHB are capable of 

sexual and asexual reproduction; however, the latter remains less costly. The degree of dispersal of 

fragments or reproductive propagules from parent sources is unknown for species along the coast but is 

likely to be highly variable. Recent research suggests dispersal distances may be greater from the 

source than predicted for many rocky intertidal species. Finally, there is essentially no information 

available on seasonal recruitment patterns for any algal species along the east Florida coast. 
Section 2 4-1 

Organismal Assemblages of East Florida Shallow Hardbottom 

Chapter 4

Invertebrates

4.1 INTRODUCTION 

There is a high diversity and abundance of sessile and motile invertebrate species associated with NHBs 

along the east Florida coast (see Appendix B, Table B.1). A total of 533 species of invertebrates has 

been identified from NHB along the east Florida coast. On a large scale, the diversity and abundance at 

any particular location varies dramatically based on latitude, depth, and, often, season. Variability in 

these community and population measures within a specific area can be high because of changes in 

features of the substrate itself, local hydrodynamics, and other biotic and abiotic factors. Often, the 

highest community biomasses occur in hardbottom areas with higher abundances of sessile invertebrate 

species (some may be considered foundational or keystone contributors to the community) that enhance 

local shelter. Higher biomasses of macroalgae that occur in the more northern hardbottoms along the 

east Florida coast likely serve the same purpose. In some cases, these hardbottom-enhancing species 

may also significantly contribute to local food webs (i.e., Phragmatopoma lapidosa). Therefore, changes 

in the abundance of these foundational species, whether natural or anthropogenic, will likely have 

profound effects on local invertebrate and vertebrate diversity and abundance. 

The approach of this chapter is to give an overview of the sessile and motile invertebrate communities 

associated with east Florida NHBs. Invertebrate community changes with latitude, depth, and season will 

be discussed, where possible, based on available data and personal research experiences. The chapter 

will additionally discuss the ecological function and life history and reproductive characteristics of each 

taxonomic group in the context of how they may affect distribution and abundance in the study region. 

4.2 SESSILE SPECIES 

4.2.1 Polychaetes 

4.2.1.1 Diversity and Trophic Function 

Diversity and abundance of polychaetes along NHBs from the east Florida coast is high (Appendix B, 

Table B.1) with 87 worm species being found along the coast. Nelson (1989) and Coastal Science 

Associates, Inc. (2000) observed 49 and 53 polychaete species, respectively, on hardbottoms in Indian 

River County, while Rudolph (1977) identified 85 species from Palm Beach County hardbottoms. 

Little is known about the ecological function of most polychaete species along the east Florida coast, but 

they are probably very important in local food webs as both consumers and prey. Most polychaetes are 

suspension feeders, although some are deposit feeders and carnivores. They are most likely very 

important food sources for various fish and invertebrate species, although no research has addressed this 

in the study region. 

Most available research on polychaetes has focused on various aspects of the reef-building polychaete 

Phragmatopoma lapidosa, also known as P. caudata (Kirtley, 1994) and/or P. l. lapidosa (Pawlik, 1988). 

This species is clearly a keystone contributor to the biological diversity of hardbottoms along the Florida 

coast. The relief provided by these "worm reefs" supports a higher diversity and abundance of marine 

species than that of neighboring sand or hardbottom habitats (Figure 4.1). In particular, worm reefs are 

considered important sources of food and shelter for juvenile green turtles (Chelonia mydas) (Wershoven 

and Wershoven, 1989; Ehrhart et al., 1996; Holloway-Adkins, 2001). Prior studies have also identified 

that the reefs provide shelter for over 423 invertebrate species (Gore et al., 1978; Nelson, 1988, 1989; 

Nelson and Demetriades, 1992) and over 200 fish species (Gilmore, 1977; Gilmore et al., 1981; 

Lindeman, 1997a; Lindeman and Snyder, 1999). The importance of worm reefs in providing food and 

shelter for juvenile and adult fishes of commercial value resulted in their designation as EFH/HAPC by the 
Section 2 4-2 

Organismal Assemblages of East Florida Shallow Hardbottom 

NMFS. In addition, worm reefs are important for the maintenance, propagation, and persistence of 

beaches and barrier islands by primarily retaining sediments (Kirtley, 1966, 1967; Multer and Milliman, 

1967; Gram, 1968; Kirtley and Tanner, 

1968; Mehta, 1973; Kirtley, 1974). 

While P. lapidosa is important in enhancing 

shelter in hardbottoms, they may also be 

important within food webs along the east 

Florida coast (Figure 4.2). It is apparent 

that various decapods (Gore et al., 1978) 

and gastropods (Watanabe, 2002; 

Watanabe and Young, 2006) will prey on P. 

lapidosa. It is likely, though not confirmed, 

that some fish species such as parrotfish 

may feed on them as well. Further, the 

rock-boring urchin Echinometra lucunter

clearly bores into worm reef and limestone, 

causing worm mortality along hardbottoms 

in St. Lucie County (D. McCarthy, 

Jacksonville University, Jacksonville, FL, 

pers. observ.). However, these biotic forms 

of mortality probably do not affect P. lapidosa abundance and distribution at the same level that seasonal 

natural disturbances such as high wave energy and sand scouring/burial (McCarthy, 2001). 

Figure 4.2. The trophic subweb postulated for the six most abundant species of decapod crustaceans on 

central Florida sabellariid worm reefs. The numbers on the lines indicate the percent of 

observed prey material noted in the gut contents from all combined individuals of a species. 

Trophic lines without numbers, or with question marks, are interpolated from our field 

observations or from records in the literature. Dashed lines indicate production from 

Phragmatopoma lapidosa (From: Gore et al., 1978). 

Figure 4.1. Intertidal worm rock at Bathtub Reef, Stuart, 

Martin County, Florida. Photo by D. McCarthy. 
Section 2 4-3 

Organismal Assemblages of East Florida Shallow Hardbottom 

4.2.1.2 Latitudinal and Cross-shelf Distribution 

There is no information currently available on how 

diversity and abundance of most polychaetes change 

with latitude and depth along east Florida coast NHBs. 

Inconsistent sampling methods for the few available 

studies make comparisons invalid. While Rudolph (1977) 

found nearly twice as many polychaetes as Nelson 

(1989), the later author suggests that increased sampling 

in Indian River County is likely to reveal more species 

than the 49 encountered in his study. Further, both of 

these studies focused only on species encountered 

within a depth of 2 to 4 m. There are no known 

comparative ecological studies of polychaete 

assemblages with increasing depth along the coast. 

The best available research for the east Florida coast has 

been conducted on Phragmatopoma lapidosa (Family 

Sabellariidae) (Figure 4.3). P. lapidosa grasps sediment 

grains from the water column by its oral tentacles, coats 

them with a proteinaceous cement, and then implants them into tubes with their opercular paleae (Kirtley, 

1966). Numerous individuals make tubes that then form vast hardbottom reefs in intertidal and shallow 

(generally highest abundances occur in <4 m) subtidal waters from Cape Canaveral to Key Biscayne in 

Florida (about 650 km) and further south to Santa Catarina, Brazil (Kirtley, 1994) (Figures 4.4 and 4.5). 

Within Florida, there appears to be higher abundances of this species towards the northern half of the 

east Florida hardbottom area (USFWS, 1999), with peak abundance possibly occurring within the Stuart 

to Vero Beach area. This may be because this region has the optimal range of sediment sizes, 

hydrodynamics, and sea water temperature. For example, juvenile worms most often use small, dark 

grains of heavy minerals, while adults tend to use larger, light-colored sand grains (Kirtley, 1966; 

Eckelbarger, 1976). In Stuart, McCarthy (2008) has observed the percentage of total intertidal worm reef 

occupying the hardbottom at Bathtub Reef to vary between <10% and >45% during his 6-year study 

(Figure 4.6). Northern limits of P. lapidosa may be controlled by the cooler sea water temperatures that 

occur towards Cape Canaveral. Eckelbarger (1976) observed significantly higher mortality (LD50) of 

P. lapidosa at temperatures below 15.5oC and above 29.5oC. Temperatures to the north of St. Lucie 

County can be as low as 7oC (Smith, 1981). To the south, McCarthy et al. (2008) suggests that 

P. lapidosa may have lower abundance in south Florida and throughout most of the Caribbean because 

of lack of availability of small sediment sizes for new recruits to build tubes. 

Figure 4.3. Close up of the anterior end of 

the reef-building polychaete 

Phragmatopoma lapidosa (caudata). 

Photo by D. McCarthy. 

Figure 4.4. Close up of the anterior tube of an 

individual Phragmatopoma lapidosa

(caudata). Photo by D. McCarthy. 

Figure 4.5. Honeycomb structure created by 

thousands of Phragmatopoma 

lapidosa (caudata). Photo by 

D. McCarthy.
Section 2 4-4 

Organismal Assemblages of East Florida Shallow Hardbottom 

Month and Year

J J ASOND J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND

% Cover

0

10

20

30

40

50

60

Live adults

Juveniles

Destroyed worms

Vacant mound

N = 10

2002 2003 2004 2005 2006 2007 2008

Figure 4.6. Monthly mean percentage cover of live adults, juveniles, destroyed worms, and vacant 

mounds for Phragmatopoma lapidosa on Bathtub Reef, Stuart, Martin County, Florida 

(McCarthy, 2008). 

4.2.1.3 Reproduction and Life History 

There is a wide range of reproductive and life history strategies for the polychaetes that have been 

identified along the east Florida coast. Most polychaetes become reproductive within a year of settlement 

and probably do not live more than 3 years (Giangrande, 1997). They are primarily known to reproduce 

sexually and have external fertilization; however, there is little known about seasonal spawning patterns 

of polychaetes in the study region. Among polychaetes in general, some spawn throughout the year, 

while others are seasonal reproducers. Regardless, most produce trochophore larvae that live in the 

plankton from weeks to months, depending on the species (Pernet et al., 2002). Consequently, larval 

dispersal varies based on species (Giangrande, 1997). Very little is known about seasonal recruitment 

patterns for most polychaetes along the east Florida coast. 

Like many polychaetes, P. lapidosa are dioecious worms that externally spawn gametes into the water, 

where fertilization results in larvae that drift in the plankton for 2 to 20 weeks (Eckelbarger, 1976; Pawlik 

and Mense, 1994). Laboratory studies (Pawlik et al., 1991; McCarthy et al., 2002) suggest that 

competent larvae position themselves deep in the water column and consequently may be more likely to 

be transported inshore during seaward-directed winds that result in near-bottom return flow. This larval 

behavior may also serve to limit dispersal among P. lapidosa populations within its rather large range. 

A molecular study by Drake et al. (2007) suggested that P. lapidosa sampled in Florida (Miami) and those 

sampled in the Caribbean (Puerto Rico and Virgin Islands) may be two distinct populations. Combined, 

these data support that P. lapidosa may have limited dispersal along the Florida coast. Future research 

is needed to establish whether there are important Florida populations that provide larvae to other Florida 

populations. 

It has been well documented that P. lapidosa require hard substrates for settlement and will repeatedly 

“test” substrates before settling on the preferred one (Eckelbarger, 1976; Pawlik, 1988). Behavioral 

preferences for moderate amounts of current have been quantified in laboratory studies with P. californica

(Pawlik et al., 1991; Pawlik and Butman, 1993). Further, it is well known that P. lapidosa often settle and 

metamorphose in response to contact with the tubes of adult worms (Eckelbarger, 1976; Krueger, 1976). 

Chromatographic analysis of the organic content of the tubes of the closely related Phragmatopoma 

californica revealed that a mixture of free fatty acids was responsible for inducing settlement (Pawlik, 

1986). Additionally, laboratory and field experiments by Jensen and Morse (1990) indicated that 

P. californica larvae also respond to butylated hydroxytoluene (BHT), which they suggest mimics the 
Section 2 4-5 

Organismal Assemblages of East Florida Shallow Hardbottom 

activity of an unidentified, cross-linked dihydroxyphenylalanine residue used to construct the tubes. 

Laboratory larval settlement responses of P. californica have been found to be identical to those of 

P. lapidosa (Pawlik, 1988). 

Recruitment occurs throughout most of the year for P. lapidosa but peaks during fall and winter 

(McCarthy, 2001). Among five intertidal locations along the Florida coast (Satellite Beach, Stuart, Coral 

Cove, Palm Beach, Boynton Beach), it appears that P. lapidosa recruit at the highest levels in Stuart, 

followed by Satellite Beach. McCarthy (2001) suggested that subtidal recruitment may be highest during 

seasons with increased wave activity. Overall, worm reefs appear to go through predictable patterns of 

annual change (McCarthy, 2001). These changes include high recruitment in early autumn through 

winter, rapid reef growth (approximately 0.5 cm/day) resulting in maximum structure in spring and 

summer (Figures 4.7 and 4.8), and decay by early autumn (Figure 4.9). McCarthy (2001) noted that 

sublethal disturbance may cue spawning from mid-winter through early fall. Based on the annual cycle of 

reef growth and decline, it seems that physical processes rather than predator/prey relationships within 

the food web are more likely to limit the abundance and diversity of flora and fauna on worm reefs 

(Applied Biology, Inc., 1979; McCarthy, 2001). When these data are integrated with those of Lindeman 

(1997a), they reveal important links between the seasonal cycle of sabellariid reef expansion and 

degradation, and the occupation of those reefs by juvenile and adult organisms. 

Figure 4.7. Phragmatopoma lapidosa (caudata) recruits recovered after 5 weeks deployment in intertidal 

hardbottom off Boynton Beach, Florida. Recruitment plates were 9 cm by 9 cm. Photo by 

D. McCarthy. 

Figure 4.8. High worm reef cover and wave activity at groynes at Palm Beach, Florida. Photo by 

D. McCarthy.
Section 2 4-6 

Organismal Assemblages of East Florida Shallow Hardbottom 

Figure 4.9. Seasonal reproductive and life history trends of Phragmatopoma lapidosa in the Boynton 

Beach area as illustrated by McCarthy (2001). 

McCarthy (2001) studied the recruitment of sessile organisms in the intertidal and subtidal (3 to 4 m) 

zones at Boynton Beach, Florida from June 1997 to January 2000. He reported 23 sessile species 

recruiting to his settlement plates, with P. lapidosa being most commonly encountered (Table 4.1). The 

recruitment rate of some 7 to 11 species m-2 d-1 occurred in both zones and was highest in summer and 

early fall. Although recruitment generally occurred at both depth zones, it was higher in the subtidal zone 

compared to the intertidal zone. McCarthy (2001) suggested that while high numbers of species may 

recruit into these habitats, most probably do not survive to adulthood, possibly due to a combination of 

factors. First, there was likely high mortality of recruits that could not tolerate frequently-occurring 

physical factors such as sand scouring and smothering. Second, high rates of P. lapidosa recruitment 

were observed to limit growth of sessile organisms, often growing over them. Finally, he also suggested 

that predation may not be as important in explaining recruit mortality in these two zones as the two 

previously-mentioned reasons. 

4.2.2 Anthozoans 

4.2.2.1 Diversity and Trophic Function 

Anthozoans are fairly abundant and diverse (54 species identified) along all NHBs from the east Florida 

coast (Appendix B, Table B.1). Most species provide shelter for various organisms via their external 

features. A number of invertebrates and fishes take refuge in the shelter created by corals. 

Consequently, when abundant, they are very important shelter-enhancing invertebrates that contribute to 

increased local biodiversity. For example, in the Florida Keys, 517 fish and 2,059 invertebrate species 

have been identified in coral reef habitats (Causey et al., 2002). Further, anthozoans such as gorgonians 

and some hard corals can serve as food sources for reef residents such as gastropods, bristle worms, 

and parrotfish (Birkland, 1974; Kinzie, 1974; Birkland and Gregory, 1975; Preston and Preston, 1975; 

Harvell and Suchanek, 1983; Lasker, 1985; Rotjan and Lewis, 2005) but are not likely to be very 

important on a large scale (O’Neal and Pawlik, 2002). 
Section 2 4-7 

Organismal Assemblages of East Florida Shallow Hardbottom Table 4.1. Species recruitment into intertidal (I) and subtidal (S) habitats from June 1997 through January 2000 in Boynton 

blue areas indicate simultaneous settlement into intertidal and subtidal habitats. Pink shading indicates habitat 

(From: McCarthy, 2001). 1997 1998 

Species Jun Jul Aug Sep Nov Jan Feb Apr Jun Jul Sep Oct Jan Mar M

Zone I S I S I S I S I S I S I S I S I S I S I S I S I S I S I 

Anomia simplex a p p p a a a p a a a a a a a p a p a p a p a p a p a a a Pinctada imbricata a p p p p p p p a p a a a a p p a p p p p p a p a p a p p Hipporina americana a p a p a a p p a a a p a p a p a p a p a p a p a p a p a Canopeum reticulum a p a a a p p p p p a p a p p p a p a a a p p p a p a p a Balanus eburneus a a a p a a p p p a a p a p a a a a a a a p a p a a a a a Chthalamus fragilis p a p a p a p a p p a a a p a a p a p p p a p a a a a p a Cliona sp. a a a a a a a a p p a p a p a p p p a p p p a p a p a a a Phragmatopoma lapidosa a a p a p a p a p a p p p p p p p a a a p p p p p p p p p Watersipora subviodea a p a p a a a a a p a p a p p p a p a p a a a p p p p p p Didemnum sp. a p a p a p a p a p a p a p a p a p a p a p a p p p a p a Diplosoma sp. a p a p a p a p p p p p p p p p p p a p a p a p p p p p a Balanus amphitrite a a a a a a p a a p a p a a a p a p a p a p a p a p a a p Brachiodontes citrinus a p a a p a a a a a a a a a a a a a a a a a a p a a a a a Phallusia nigra a p a a a a a p p a a a a a a a a a a a a a a a a a a a a Botryllus sp. a a a a a a p p a a a a a a a p a a a a a a a a a a a a a Bugula sp. a a a a a a a a a a a a a a a a a a a p a a a a a a a a a Hydroides dianthus a p a p a a a a a a a p a a a p a a a p a p a p a a a a a Terebellid sp. a a a a a a a a a a a a a a a a a a a a a a a a a a a a a Actinia bermudensis a p a a a a a a p p a a a a a a p a a a a a a a a a a a a Siderastrea radians a a a a a a a a a a a a a a a a a a a a a p a a a a a a a Unknown blue sponge a a a a a a a a a a a a a a a a a a a a a p a a a a a a a Vermicularia sp. a a a a a a a a a a a a a a p a a a a a a a a a a a a a a Turbicellaora avicularis a a a a a a a a a a a a a p a a a p a p a a a p a a a 

Species number at each habitat 1 11 4 8 4 4 8 9 8 9 2 10 2 9 6 13 5 9 2 12 4 14 3 14 4 11 3 9 4 Species number at both 0 2 1 5 5 2 2 5 2 2 3 2 4 3 0 Total see number 12 10 8 12 12 10 9 14 12 12 15 15 11 9 4 

a = absent; p = present. Subtidal species Cthalamus fragilis was more intertidal than subtitled and experienced significant, predation; Balanus eburneus recruited to both intertidal and subti

(Vermicularia sp.) and intertidal.
Section 2 4-8 

Organismal Assemblages of East Florida Shallow Hardbottom 

4.2.2.2 Latitudinal and Cross-shelf Distribution 

The diversity and abundance of anthozoans changes dramatically with latitude, and to a lesser extent 

with depth, along the east Florida coast NHBs. Throughout tidal pools in the intertidal and very shallow 

subtidal zones along the entire coast, there are few of 

these animals present. The species that can be 

encountered include the starlet coral Siderastrea spp.

(Figure 4.10), two species of zoanthids (Palythoa 

caribaeorum and Zoanthus pulchellus) (Figure 4.11), 

and several species of solitary anemones 

(Bunodosoma caveranta, Actinia bermudnesis, 

Diadumene leucolena). These are species that 

generally have a higher tolerance for the fluctuations 

in salinity and temperature that occur in these 

habitats (Muthiga and Szmant, 1987; Lirman et al., 

2002). P. caribaeorum is also known to be a very 

fast growing highly aggressive competitor for space 

among coral species (Suchanek and Green, 1981). 

Progressing into the shallow subtidal area these 

species probably are more abundant, although there 

are no available data to support this statement. 

Generally, anthozoans are not very abundant within 

the 0 to 4 m depth range. 

Anthozoan diversity in the subtidal 

hardbottom changes most significantly with 

latitude. However, there are very few 

studies of anthozoan communities in water 

depths less than 4 m. North of Martin 

County, there are generally fewer hard 

corals. Besides small Siderastrea spp., the 

most frequently encountered hard corals are 

the Oculina species (O. varicosa and O. 

diffusa). The latter two species become 

fairly abundant in water depths in excess of 

6 m (Reed et al., 1982; D. McCarthy, 

Jacksonville University, Jacksonville, FL, 

pers. observ.). However, there are 

occasional occurrences of Cladocora 

arbuscula and grouped polyps of Phyllangia

americana and Astrangia spp. (Brooke and 

Young, 2005). It is in the deeper NHB that 

Oculina spp. are likely most important in 

contributing significant shelter for 

invertebrate and vertebrate species (Reed, 

1982; Reed et al., 1982; Reed and 

Mikkelsen, 1987). The shelter value of the 

much deeper (70 to 152 m) O. varicosa reefs 

has been recognized and has resulted in their being listed as an HAPC by the SAFMC (NMFS, 2007). In 

terms of soft corals within NHB, two species of gorgonians (Leptogorgia virgulata and Leptogorgia hebes) 

and one species of telestacean (Carijoa [Telesto] riisei) are fairly common in the 4 to 6 m depth range. 

Progressing southward, diversity and abundance of anthozoans, particularly scleractinians, increase on 

subtidal hardbottoms (Southeast Florida Coral Reef Evaluation and Monitoring Project [SFCREMP], 

2007; J. Beal, Florida Fish and Wildlife Conservation Commission [FWC], pers. comm.). In Martin 

County, the St. Lucie Inlet Reef appears to be the northernmost extent of several of the reef-building 

Figure 4.11. The zooanthid Zoanthus pulchellus and 

various other sessile invertebrates (and green 

algae) on intertidal hardbottom in Palm 

Beach, Florida. Photo by D. McCarthy.

Figure 4.10. An intertidal colony of the 

scleractinian Siderastrea sp. in 

Palm Beach, Florida. Photo by 

D. McCarthy.
Section 2 4-9 

Organismal Assemblages of East Florida Shallow Hardbottom 

corals (i.e., Diploria and Montastraea spp.) that are known important biodiversity contributors within the 

Caribbean region. Additionally, a high number of octocoral species (~15 spp; J. Beal, FWC, pers. comm.) 

are encountered at this latitude. Consequently, this reef was recently designated for protection as part of 

the St. Lucie Inlet Preserve State Park because of the diversity and abundance of corals and other 

shelter-enhancing invertebrate species (polychaetes, sponges, tunicates, algae) present. 

Both diversity and abundance of stony coral species on NHB increase southward from Martin County 

(Appendix B, Table B.1; SFCREMP, 2007). A synthesis of available reports revealed that 7, 22, 24, and 

32 stony coral species were identified in NHB in St. Lucie, Martin, Palm Beach, and Broward Counties, 

respectively. The SFCREMP (which encompasses sites in Martin, Palm Beach, Broward, and Dade 

Counties) also documented an increase in stony coral diversity towards Dade County, although observed 

species numbers are low because of the scientific approach used. In their study, the highest diversity of 

corals among shallower sites (17 to 19 species) occurred in Dade and Broward Counties compared to 

Palm Beach and Martin Counties (8 to 9 species) (SFCREMP, 2007). They also observed that seven 

species of stony corals (Dichocoenia stokesii, Diploria clivosa, Millepora alcicornis, Montastraea 

cavernosa, Porites astreoides, Siderastrea siderea, and Solenastrea bournoni) were present in all four 

counties from Martin to Dade County (SECREMP, 2007). However, it should be noted that in order to 

compare all four locations, only the shallowest sites (8 m in Palm Beach, Broward, and Dade; 5 m in 

Martin County) were used, which is outside the designated NHB depth range. In shallower hard bottoms 

(<7 m), Prekel at al. (2007, 2008) and Coastal Planning & Engineering, Inc. (2006) recorded 19 stony 

coral species on NHB in Broward County, observing that Siderastrea siderea was most common 

(81.3% of observations of corals), followed by Porites porites, Dichocoenia stokesii, Phyllangia 

americana, and Porites astreoides (accounting for between 1.1% to 4.9% of observations). Also, in 

Broward County, hardbottom areas have been observed at the 4 m depth and deeper, with fairly high 

percent cover of the scleractinian Acropora cervicornis (Moyer et al., 2003; Vargas-Angel et al., 2003; 

SFCREMP, 2007). The tree-like morphology of this coral offers important shelter value to motile 

invertebrates and vertebrates. Consequently, in 2006 this species (along with A. palmata) was 

designated by the National Oceanic and Atmospheric Administration (NOAA) as threatened under the 

ESA because of its significant decline and subsequent risk of extinction within this Florida-Caribbean 

region. 

The diversity and abundance of octocoral species on NHB likely increases southward from St. Lucie 

County, although there is not a great amount of data that clearly identifies whether species were 

observed within NHB or deeper (Appendix B, Table B.1). In this study, a synthesis of available reports 

revealed that 3, 12, 14, and 14 octocoral species were identified in NHB in St. Lucie, Martin, Palm Beach, 

and Broward Counties, respectively. However, over 40 octocoral species have been identified in hard 

bottom habitats up to 8 m in depth from Palm Beach to Dade Counties (Jaap, 1984; Vare, 1991). 

4.2.2.3 Reproduction and Life History 

Sexual vs. Asexual Reproduction 

There is high diversity in reproductive methods among anthozoan species found along east Florida. Most 

species can reproduce and/or propagate both sexually and asexually (see Fautin, 2002 for a review of 

cnidarian reproduction). However, the relative importance of these forms of reproduction in maintaining 

population numbers varies with the species and sometimes is unclear. Asexual reproduction via 

longitudinal fission appears to be the dominant form of vegetative proliferation among sea anemones 

(Shick, 1991). This may not be important for most colonial scleractinians relative to sexual reproduction, 

but it has been suggested to be the dominant form of reproduction for Acropora cervicornis, whereby 

propagation occurs when branches break off during a storm and new colonies form (NOAA Fisheries, 

2008). There is less information available on gorgonians, but it has been suggested that Plexaura may 

reproduce via parthenogenesis, whereby eggs develop without the aid of fertilization (Brazeau and 

Lasker, 1989). This species may also asexually reproduce when branches break during storms (Lasker, 

1984). 
Section 2 4-10 

Organismal Assemblages of East Florida Shallow Hardbottom 

Seasonal Spawning Patterns 

There have not been many studies documenting seasonal reproductive patterns for anthozoans along the 

Florida coast, but it is likely that many species spawn at the same times as their Caribbean counterparts. 

Some species are known to externally fertilize, while others brood their larvae. A number of the 

scleractinian species in the region have been noted to spawn during late summer. To the north, Oculina

spp. spawns during late summer, producing planula that can settle in 3 weeks (Brooke and Young, 2005). 

While not confirmed for Florida, Leptogorgia spp. and Pseudoplexaura porosa may spawn seasonally 

during this time period as well (Beasley et al., 1997; Kapela and Lasker, 1999). In southeast Florida, the 

reef building corals Montastraea annularis, M. cavernosa, Acropora cervicornis, and A. palmata

(Szmant-Froelich, 1986; Vargas-Angel et al., 2006) also spawn during this time period. In Martin County, 

it has been suggested that Montastraea and Diploria colonies do not spawn because they are at the 

northern limit of their distribution and do not have the energetic reserves to produce gametes (J. Beal, 

FWC, pers. comm.). 

In contrast to seasonal spawning, some anthozoans may spawn for most of the year. Siderastrea radians

and Porites astreoides brood larvae during an extended season and reach the ability to reproduce at 

smaller sizes than species such as Diploria spp. and M. cavernosa (Soong, 1991, 1993). S. radians may 

be encountered throughout the east Florida coast in intertidal and shallow subtidal hardbottoms because 

of its strong tolerances to environmental fluctuations and continuous reproductive capability, which allow 

for it to be able to take advantage of available space created by natural and anthropogenic disturbances 

throughout the year. 

Larval Transport and Recruitment 

Little is known about larval transport and recruitment for anthozoans along the east Florida coast. The 

amount of time that larvae can be in the plankton varies from days to months. Some species of zoanthids 

have larvae that are well-known for being far dispersed in the plankton. However, while they may survive 

there for as long as 22 weeks, their ability to metamorphose may be lost after 11 weeks (Ryland, 1997). 

Brooke (2002) suggests that larvae of the ivory bush coral (Oculina varicosa) not only have the potential 

to be transported between the deep reef tracts but may also contribute to nearshore shallow water 

populations during summer upwelling events. In contrast, the staghorn coral Acropora cervicornis likely 

has low dispersal and hence gene flow, which suggests that this endangered species require local source 

populations for their recovery (Vollmer and Palumbi, 2007). 

Generally, the few studies of hardbottom recruitment along the east Florida coast have observed 

infrequent and sporadic seasonal recruitment of anthozoans. Yoshioka (1996) studied recruitment of 

several gorgonian species on the Atlantic coast of Panama, observing limited yet highly variable seasonal 

recruitment. Both Cummings (1994) and McCarthy (2001) observed seasonal recruitment of sessile 

invertebrates to hardbottoms (4 to 8 m depths) in southeast Florida and encountered only one 

scleractinian species (S. radians). McCarthy (2001) sampled plates frequently in Boynton Beach for 

~2.5 years, encountering this species only once out of over 1,200 settlement plates sampled. Cummings 

(1994) additionally observed the anthozoans Carijoa (Telesto) riisei, Bunodosoma granuliferum, and 

Palythoa mamillosa, while McCarthy encountered only the actiniarian Actinia bermudensis. 

Consequently, it appears that anthozoan recruitment is low in these habitats and may not provide rapid 

recovery and/or maintenance of coral populations after natural or anthropogenic disturbance events. 

Such recovery may be further reduced by cyanobacteria and macroalgal blooms, which can prohibit coral 

recruitment and appear to be more frequent along the east Florida coast in recent years (Paul et al., 

2005; Ritson-Williams et al., 2005; Kuffner et al., 2006). 
Section 2 4-11 

Organismal Assemblages of East Florida Shallow Hardbottom 

4.2.3 Sponges 

4.2.3.1 Diversity and Trophic Function 

Sponges are fairly abundant and diverse (50 species identified) on all NHBs along the east Florida coast 

(Appendix B, Table B.1). Most species provide shelter for various organisms via internal spaces as well 

as their external features. Amphipods, crabs, shrimp, nemertines, polychaetes, molluscs, and brittle stars 

all take refuge in sponges (Wulff, 2006; D. McCarthy, Jacksonville University, Jacksonville, FL, pers. 

observ.). Consequently, when abundant, sponges are important shelter-enhancing invertebrates that can 

contribute to increased local biodiversity. Sponges may also serve as an important food source for many 

of the angelfishes, filefishes, trunkfishes, cowfishes, and butterflyfishes that are particularly common in 

southeast Florida (Pawlik, 1998). As filter feeders, they also may be important in removing nutrients 

(e.g., carbon, nitrogen, oxygen) from the water column and thus significantly impact pelagic ecosystems 

(Peterson et al., 2007; see Bell, 2008 for a review). Finally, some sponges can have a detrimental effect. 

For instance, the rock-boring pionid (formerly clionid) species can aid in the fragmentation and 

disintegration of coral heads (Glynn, 1997; Zilberberg et al., 2006). The rate of this type of coral 

disturbance may be accelerated by heavy storms, hurricanes, and any other source of physical stress 

(Tunnicliffe, 1981; Highsmith, 1982; Rutzler, 2002). Such bioerosion by sponges results in the production 

of reef sediment, which can affect the structural integrity of coral (Rutzler, 1975; Glynn, 1997). Further, 

the proliferation of clionid sponges has been linked to organic pollution and temperature increases (Rose 

and Risk, 1985; Holmes, 1997; Rutzler, 2002). For example, Ward-Paige et al. (2005) suggested that 

sewage contamination in the Florida reef tract results in increases in bioeroding sponges that shift the 

carbonate balance from one of reef construction (accretion) to that of reef destruction (erosion). 

4.2.3.2 Latitudinal and Cross-shelf Distribution 

The diversity of sponges probably 

increases with latitude and depth along 

the NHBs along the east Florida coast. 

However, abundance patterns are 

unclear for most species, with little 

quantitative information being available 

for the 0 to 4 m depth range. Within the 

5 to 8 m depth range, mean percent 

cover of sponges is highly variable, 

ranging between 0.14% and 10.29% in 

the survey areas (SFCREMP, 2007). 

Within the lower intertidal and shallow 

subtidal zones, the most conspicuous 

sponge is the rock-boring sponge Pione 

lampa (formerly Cliona lampa) 

(Figure 4.12). It probably continues on 

to deeper depths where other rock 

boring sponges such as Pione delitrix

can be encountered. SFCREMP (2008) 

observed P. delitrix at the 5 to 8 m depth 

range for the four locations it studied 

along the east Florida coast. There were no latitudinal trends with this species, with total colony areas 

varying between 3.2 and 32.16 cm2

/m2. 

4.2.3.3 Reproduction and Life History 

Sponges can reproduce sexually and asexually, although the relative importance of these strategies in 

population maintenance is often unclear. Asexual reproduction (i.e., fragmentation, budding) may be 

important for a number of the shallower sponge species because of the high frequency of lethal and 

sublethal disturbances that result from high wave energy and sand scouring. Some more 

Figure 4.12. An intertidal colony of the rock-boring sponge 

Pione (Cliona) lampa in Stuart, Florida. 

Photo by D. McCarthy.
Section 2 4-12 

Organismal Assemblages of East Florida Shallow Hardbottom 

vertically-oriented sponges common off east 

Florida such as Aplysina spp. (Figure 4.13) and 

Iotrochota birotulata may propagate exclusively by 

asexual fragmentation (Wulff, 1991). Schönberg 

(2002) suggests that P. lampa can produce 

gemmules (normally found in freshwater species) 

that can resist sand burial, heat, and desiccation 

and ensure survival in these harsh habitats. 

Additionally, gemmules may be scattered and 

dispersed during periods of increased wave 

activity along the coast. 

Sexually, fertilization in most sponges is internal, 

with embryos being either brooded or shed 

immediately into the water column. In both cases, 

the resultant larvae either swim in the water 

column or crawl away on the substrate 

(Maldonado and Bergquist, 2002). For species 

with planktonic larvae, most appear to be 

short-lived lecithotrophs living in the plankton from 

a few hours to several days (Lindquist et al., 1997; 

Bergquist, 2002). 

There is very little information on seasonal spawning, dispersal, and recruitment patterns of sponges 

along the Florida coast. Cummings (1994) often encountered the rock-boring sponge Pione (formerly 

Cliona) lampa during her recruitment study in 2 m of water off Boca Raton. She less frequently 

encountered Chondrilla nucula, Spongia sp., Demapsamma sp., and one unknown species. 

4.2.4 Hydrozoans 

4.2.4.1 Diversity and Trophic Function 

Various hydrozoans can be abundant along all NHBs along the east Florida coast but are generally lower 

in diversity (12 species identified) compared to some of the other sessile invertebrates discussed 

(Appendix B, Table B.1; Figure 4.14). Some species may provide limited shelter for some small 

organisms, but most probably do not. With their relatively small biomass and nematocyst defense 

capability, it is unlikely, that they are an important part of the food web, although this is not known. 

Figure 4.14. Colonial hydrozoans on a mitigation reef in Juno Beach, Florida. Photo by D. McCarthy. 

Figure 4.13. The demosponge Aplysina fistularia on 

hardbottom off Palm Beach, Florida. Photo 

by D. McCarthy.
Section 2 4-13 

Organismal Assemblages of East Florida Shallow Hardbottom 

4.2.4.2 Latitudinal and Cross-shelf Distribution 

It is unknown how the diversity and abundance of hydrozoans change with latitude and depth along east 

Florida coast NHBs. SFCREMP (2007) encountered the fire coral Millepora alcicornis at a depth of 5 m at 

the St. Lucie Reef but not at any of the other shallower water sites (all other sites were ~8 m) in southeast 

Florida. However, M. alcicornis was fairly abundant at deeper sites surveyed. There are few known 

hydroid species within the intertidal zone, but generally several that are common within the shallow 

subtidal zone. Some of the more commonly occurring hydroid species along the coast include 

Thyroscyphus ramosus, Campanularia sp., Eudendrium ramosum, Lytocarpus philipinus, Pennaria sp., 

and Sertularella sp. 

4.2.4.3 Reproduction and Life History 

Hydrozoans are primarily colonial organisms that have life cycles that may include polyps, medusae, or 

both. Reproduction can occur via asexual and/or sexual reproduction. Generally, clonal reproduction by 

budding is most common among the solitary A-form polyps (Ruppert et al., 2004). Offspring that may drift 

in the plankton (or creep along the sea floor bottom) include asexually produced medusae or planula 

larvae that are formed via sexual reproduction (external or internal fertilization) of medusae or polyps 

(Ruppert et al., 2004). Hydrozoan larvae are generally short-lived in the plankton. For instance, 

Mitrocomella polydiademata are ready to settle 3 to 5 days after fertilization (Martin and Koss, 2002). 

Consequently, there probably is very limited dispersal among offspring from their parents. 

There is very little information on seasonal spawning and recruitment patterns of hydrozoans along the 

Florida coast. If local species are like temperate hydrozoans, they may reproduce asexually throughout 

the year but sexually on a seasonal basis (Coma et al., 1996). Cummings (1994) observed recruitment of 

Thyroscyphus marginatus, Eudendrium spp., Lytocarpus philipinus, Pennaria sp., and Gymnangium spp. 

throughout the year. In contrast, Walton Smith et al. (1950) observed very low recruitment of hydroids in 

the Key Biscayne area. The most common identifiable hydroid in that study was Tubularia sp. 

4.2.5 Tunicates and Bryozoans 

4.2.5.1 Diversity and Trophic Function 

Tunicates and bryozoans are fairly abundant and diverse (93 bryozoan species) along all NHBs along the 

east Florida coast (Appendix B, Table B.1). Most occur in colonies (branching or encrusting forms), 

although there are some solitary forms of tunicates that are abundant (i.e., Phallusia, Mogula). Both 

groups are particularly abundant under ledges and within crevices. Most species probably do not provide 

very large shelter enhancement to the habitat, but some do provide shelter to smaller species such as 

amphipods, crabs, shrimp, nemertines, polychaetes, molluscs, and brittle stars that may take refuge 

along the edge of colonies or within some branching forms (i.e., Bugula spp.) (Voultsiadou et al., 2007; 

D. McCarthy, Jacksonville University, Jacksonville, FL, pers. observ.). It is unlikely that many of the 

tunicates are important in the diet for any hardbottom predator, as many produce secondary metabolites 

that deter at least fish predators (Pisut and Pawlik, 2002; Odate and Pawlik, 2007). Bryozoans may be 

more susceptible to predation, although they have morphological defenses to deter predators as well 

(Harvell, 1992; Ruppert et al., 2004). 

4.2.5.2 Latitudinal and Cross-shelf Distribution 

There is not a great deal of information available on how the diversity and abundance of tunicates and 

bryozoans change with latitude and depth along the east Florida coast NHBs. The most conspicuous 

animals of these types found in tide pools of the lower intertidal zones and shallow subtidal habitats are 

the colonial tunicates Botryllus planus (Figure 4.15), Botrylloides nigrum, Didemnum spp. (probably most 

common), and the bryozoan Watersipora subvoidea. Progressing into the shallow subtidal area, diversity 

may slightly increase but abundance appears to remain fairly low. Essentially all bryozoans remain within 

crevices and under ledges, while several species of tunicate can be found on top of hardbottoms. Some 

common tunicate species encountered subtidally include Eudistoma sp., Diplostoma sp., Distaplia sp., 
Section 2 4-14 

Organismal Assemblages of East Florida Shallow Hardbottom 

Mogula sp., and Phallusia (Ascidia) niagra, with the 

latter species clearly more abundant from St. Lucie 

southward (D. McCarthy, Jacksonville University, 

Jacksonville, FL, pers. observ.). 

4.2.5.3 Reproduction and Life History 

Most tunicates and bryozoans along the east Florida 

coast have colonial forms and can reproduce 

asexually and sexually. Asexually, most tunicates 

and bryozoans proliferate via budding (Ruppert et al., 

2004). In the case of bryozoans, their success at 

occupying space may be largely related to their 

capacity for colonial growth via budding (Seed and 

Hughes, 1992), and this may be the case for 

tunicates as well. Sexually, individuals in both 

groups are hermaphrodites that, depending on the 

species, fertilize internally or externally. While a few 

families of tunicates have direct development, in most cases lecithotrophic larvae are produced that live in 

the plankton for a few minutes to a few hours (Clooney et al., 2002). Consequently, their dispersal 

distance from parent populations should not be very great. Bryozoan larvae may be either planktotrophic 

or lecithotrophic, but also have a very short planktonic period (Temkin and Zimmer, 2002). 

There is very little available information on seasonal spawning and recruitment patterns of tunicates and 

bryozoans in the study region. In Boynton Beach, the colonial ascidians Didemnum sp. and Diplosoma

sp. and the bryozoans Canopeum reticulum and Watersipora subzoidea appear to recruit throughout 

most of the year (McCarthy, 2001). In the same study, Bugula sp., Phallusia niagra, and Botryllus sp. 

sporadically recruited just a few times during the 2.8-year study. 

4.2.6 Molluscs (Sessile)

4.2.6.1 Diversity and Trophic Function 

A number of mollusc species occur on NHBs along the east Florida coast (Tables 4.1 to 4.4; 

Appendix B, Table B.1). The sessile forms include primarily bivalves but also two species of vermetid 

gastropods. Most bivalves are found under ledges and within crevices in somewhat low abundance. The 

vermetid gastropods occur primarily in the upper intertidal zone (D. McCarthy, Jacksonville University, 

Jacksonville, FL, pers. observ.). Both groups of molluscs can contribute to enhancing shelter on 

hardbottoms and may benefit generally smaller invertebrates (i.e., sipunculans, xanthid crabs, shrimp) 

and possibly a few small or juvenile fish species. They are likely preyed on by invertebrates and fishes, 

but no data are available on their trophic importance along the east Florida coast. 

4.2.6.2 Latitudinal and Cross-shelf Distribution 

It is unclear how the diversity and abundance of molluscs change with latitude and depth along the NHBs 

along the east Florida coast. The most conspicuous molluscs are found in the upper intertidal zones. 

These are the vermetid gastropod Petaloconchus nigricans and P. varians, which can form vast networks 

of tubes (10 to 15 cm thickness) that provide significant shelter for a number of invertebrates (xanthid 

crabs, sipunculans, polychaetes, and nemertines) and fishes (blennies, gobies, toadfishes). Eight 

species of bivalves occur at the intertidal worm reef at Stuart: Barbatia dominensis, Isognomon radiatus, 

Musculus lateralis, Ostrea equestris, Sphenia antillensis, Anomia sp., Pteria sp., and Pinctada imbricata

(McCarthy, 2008). To the north in St. Lucie County, Reed and Mikkelsen (1987) identified 40 bivalve 

species associated with the scleractinian coral Oculina varicosa at 5.5 m depth. Cummings (1994) and 

McCarthy (2001) combined identified Isognomon radiatus, Ostrea equestris, Anomia simplex, Pteria sp., 

Brachiodontes citrinus, and Pinctada imbricata on settlement plates in the 4 to 8 m depth range. 

Figure 4.15. Botryllus sp. overgrowing dead worm 

rock at Boynton Beach, Florida. Photo 

by D. McCarthy.
Section 2 4-15 

Organismal Assemblages of East Florida Shallow Hardbottom 

Table 4.2. Abundance of associated macroinvertebrates collected from sabellariid worm reef 

(Stations 9 and 10 combined) at the St. Lucie Power Plant from April 1976 through April 1979 

(From: Applied Biology, Inc., 1979). 

1976 1977 1978 1979 Species Apr Jul Oct Jan Apr Jul Oct Jan Apr Jul Oct Jan Apr 

MOLLUSCA 

Gastropoda 

Anachis avara 75 5 3 1 

Anachis floridana 2 15 16 1 11 

Anachis lafresnayi 1 1 

Astraea tuber 1 1 

Barleeia tincta 26 1 

Cypraea sp. 1 

Erato maugeriae 1 

Fissurella barbadensis 1 2 

Lamellaria perspicua 1 2 

Lucapinella Iimatula 2 

Mitrella argus 4 36 9 5 

Nudibranchia sp. 1 

Phidiana ?Iynceus 1 2 

Pisania tinctus 1 1 

Thais haemastoma floridana 1 1 

Thais rustica 3 

Tricolia affinis pterocladica 1 

Bivalvia 

Barbatia domingensis 1 

Isognomon radiatus 1 

Musculus lateralis 1 

Ostrea equestris 4 

Sphenia antiIIensis 1 1 1 1 

ARTHROPODA 

Pycnogonida 

Tanystylum sp. 2 

Stomatopoda 

Gonodactylus bredini 1 1 

Gonodactylus sp. 

lsopoda 

Accalathura crenulata 1 

Anthuridae sp. 1 

Asellota sp. 1 

Dynamenella sp. 1 1 1 2 

Excorallana sexticornis 11 5 10 51 9 14 5 56 4 42 

Mesanthura decorata 1 4 2 2 3 6 

Paranthura infundibulata 1 1 2 1 1 5 

Sphaeroma walkeri 6 1 27 70 12 1 3 4 41 7 18 
Section 2 4-16 

Organismal Assemblages of East Florida Shallow Hardbottom 

Table 4.3. Molluscs recorded during a survey of a newly constructed artificial reef structure in the 

nearshore (~8 m depth) area off Boca Raton, Florida (Adapted from: Cummings, 1994). 

● denotes species presence. 

Scientific Name December 1988-

January 1989 

July 1989- 

August 1989 July 1990 

Class Gastropoda 

Order Neogastropoda 

Anachis floridana ● ●

Thais rustica ● ● ●

Iselica fenestrata ● 

Class Bivalvia 

Order Pterioida 

Isognomon radiatus ● ● ●

Ostrea equestris ● ● 

Ostrea sp. ●

Pinctada imbricata ● 

Table 4.4. Data summary for molluscs collected from 41 colonies of the scleractinian coral Oculina 

varicosa on four reefs off the east Florida coast (From: Reed and Mikkelsen, 1987). Numbers 

in parentheses = total species. Density = number of individuals/100 g coral dry weight. 

Skeletal volume determined by water displacement. Percent dead coral based on dry weight. 

Water Depth 

6 m 27 m 42 m 80 m 

Molluscs 

Number of individuals 2,027 61 594 2,450 

Average number of individuals/coral 145 15 46 245 

Average density 29.3 9.1 3.8 14.7 

Gastropod species (155) 59 9 67 89 

Bivalve species (68) 40 15 24 46 

Total species (230) 101 25 91 140 

Mean number of species/coral 18 9 18 42 

H' Diversity 2.97 4.24 5.13 4.79 

H (Brillouin) 2.87 3.55 4.81 4.66 

E (Equitability) 0.44 0.92 0.79 0.67 

DI (Dominance) 54.4 13.1 15.8 23.9 

Corals 

Number of coral samples 14 4 13 10 

Colony dry weight (g) 196-1,207 147-204 526-2,715 279-2,658 

Skeletal volume (mL) 97-723 53-110 260-1,431 149-1,544 

Percent colony dead 0-62.8 5.7-40.0 3.4-43.2 27.5-87.9 

4.2.6.3 Reproduction and Life History 

Vermetid gastropods and bivalves are sexually reproducing invertebrates. Vermetid gastropods are 

protandrous hermaphrodites that internally fertilize, producing lecithotrophic larvae with a short, 

free-swimming/crawling phase. Bivalves usually have separate sexes and either spawn externally or 

brood larvae. Their embryos may end up in the plankton, while others may directly develop (Zardus and 

Martel, 2002). However, most bivalves produce veliger larvae that swim in the plankton for 3 to 4 weeks. 

Depending on the species, larvae can be planktotrophic or lecithotrophic. 
Section 2 4-17 

Organismal Assemblages of East Florida Shallow Hardbottom 

Little is known on seasonal spawning and recruitment patterns of these species along the Florida coast. 

In Boynton Beach, Vermicularia sp. and Brachiodontes sp. recruitment appears highly sporadic 

(McCarthy, 2001). In the same area, Pinctada imbricata recruited seasonally, with peak recruitment 

occurring during the fall (McCarthy, 2001). Larvae from bivalves generally are longer-lived in the plankton 

than many invertebrates and may have somewhat longer dispersal among offspring from their parents. 

4.2.7 Barnacles

4.2.7.1 Diversity and Trophic Function 

Barnacles are encountered throughout intertidal, and to a lesser extent, subtidal hardbottoms along the 

east Florida coast. There are nine species found along the coast (Appendix B, Table B.1). They are 

active suspension feeders that use cirri to move water and feed. When in large numbers, they can 

provide hardbottom shelter enhancement (usually in the upper intertidal zone) and serve as a food source 

for some fishes (i.e., sheepshead) and some larger decapods (stone crabs) and gastropods (oyster drills). 

However, they are not likely to be very important on a large scale due to their generally low abundance. 

4.2.7.2 Latitudinal and Cross-shelf Distribution 

There is not a great deal of information on how diversity and abundance of barnacles change with latitude 

and depth along the east Florida coast NHBs. The most conspicuous barnacle, the volcano barnacle 

(Tetraclita squamosa stalactifera), is very prominent along the coast at the upper intertidal zone, where 

the competitive P. lapidosa does not often 

occur (D. McCarthy, Jacksonville University, 

Jacksonville, FL, pers. observ.) (Figure 4.16). 

Multer and Milliman (1967) also suggested 

that T. s. stalactifera compete for space with 

P. lapidosa in this zone. McCarthy (2001) 

suggests that P. lapidosa consistently recruits 

in large numbers and out-competes other 

recruit species such as barnacles by growing 

over them. When P. lapidosa is not present, 

a number of other barnacle species may often 

be present in these habitats. The barnacle 

Balanus tintinnabulum antillensis was 

observed attached to cocinoid rock underlying 

the worm-built structure on Hutchinson Island 

(Camp et al., 1977) and on an artificial reef in 

Boca Raton (Cummings, 1994). The invasive 

barnacle Megabalanus cocopoma was 

identified on intertidal hard bottom in Stuart 

(McCarthy, 2006). McCarthy (2001) observed 

the following barnacle species on intertidal and 

subtidal hard bottoms in Boynton Beach: 

Balanus eburneus, B. amphitrite, and 

Chthamalus fragilis. Further to the south in the Key Biscayne area, additional barnacles encountered are 

Balanus improvisus and Chthamalus stellatus (Walton Smith et al., 1950). Considering relatively high 

dispersal ability, most of these species probably occur throughout intertidal and shallow subtidal 

hardbottoms along the east Florida coast. 

4.2.7.3 Reproduction and Life History 

Most barnacles are simultaneous hermaphrodites that externally fertilize eggs within their mantle cavities. 

Eggs are brooded there, hatch out into the plankton, and become nauplius larvae that are carried with 

currents until they settle as cyprids. Many cyprids may preferentially settle on conspecifics, cuing in on 

chemical and topographical features of the adults (Ruppert et al., 2004). Generally, there is perceived to 

Figure 4.16. The volcano barnacle Tetraclita squamosa

stalactifera and various other sessile 

invertebrates on intertidal hardbottom in 

Palm Beach, Florida. Photo by D. McCarthy.
Section 2 4-18 

Organismal Assemblages of East Florida Shallow Hardbottom 

be high initial post-settlement mortality, with surviving barnacles probably living 1 to 10 years, depending 

on various abiotic and biotic factors (Ruppert et al., 2004). 

Little is known about seasonal reproductive patterns of most barnacles along the east Florida coast, 

although some species may have similar patterns as counterparts within the region. McCarthy (2001) 

observed barnacle recruitment on intertidal and subtidal (4 to 5 m) hardbottom habitats in Boynton Beach. 

In the intertidal zone, all three barnacle species encountered (Balanus eburneus, B. amphitrite, and 

Chthamalus fragilis) recruited seasonally, with peak recruitment occurring during the summer time period 

(May to August). C. fragilis recruited more intertidally rather than subtidally and generally was the most 

abundant barnacle recruiter during the study. B. eburneus was the only barnacle species that recruited 

equally to both intertidal and subtidal habitats. In contrast, B. amphitrite was the most abundant 

subtidally, recruiting throughout the year, while B. eburneus and C. fragilis had sporadic recruitment 

patterns. The study also suggested that there may be significant post-settlement mortality by mobile 

predators (i.e., crabs and fishes) for C. fragilis in subtidal habitats but not for any of the other barnacle 

species. In southeast Florida, B. amphitrite and B. improvisus had similar seasonal peaks in recruitment 

as those found by McCarthy (2001) but were also found to recruit more continuously throughout the year 

(Walton Smith et al., 1950). Interestingly, Tetraclita spp. were essentially non-existent in both studies yet 

are often the most abundant adult barnacles along the coast. A possible explanation for this may be that 

settlement plates in both studies were not placed high enough in the intertidal zone to observe 

recruitment of this species. Larval behavioral preferences may have positioned such competent larvae so 

that they were transported only to these upper reaches of the intertidal zone where most adults are found. 

4.3 MOTILE SPECIES 

4.3.1 Crustaceans 

4.3.1.1 Diversity and Trophic Function 

The structure provided by east Florida coast NHBs supports a high diversity and abundance of crabs, 

stomatopods, shrimp, lobsters, isopods, and amphipods (Appendix B, Table B.1). Sabellariid reefs, in 

particular, support an even higher diversity and abundance of these crustaceans than neighboring sand 

or hardbottom habitats. Van Montfrans (1981) reported that the majority of decapods along the east 

Florida coast can be regarded as tropical in origin, 

as can be seen by their associations with southern 

latitude coral reefs. Furthermore, Phragmatopoma

worm rock reefs are an important habitat for 

tropical marine decapods (Figure 4.17), especially 

in the high energy surf zone, where corals tend not 

to prevail at the more northern latitudes. 

Trophically, crustaceans are important in food 

webs along the east Florida coast as major prey 

for other invertebrates and fishes. They can also 

be important as predators of various invertebrates 

and small fishes (USFWS, 1999). Gore et al.

(1978) postulated a possible food sub-web for 

sabellariid colonies on the east Florida coast 

based on the gut contents of six common species 

collected. The worms form the basis of the food 

sub-web on which some species of reef-dwelling 

crabs forage (Figure 4.2). 

Figure 4.17. Tidal spray crab (Plagiusa depressa) 

in Palm Beach, Florida. Photo by 

D. McCarthy.
Section 2 4-19 

Organismal Assemblages of East Florida Shallow Hardbottom 

4.3.1.2 Latitudinal and Cross-shelf Distribution 

The most quantitative research with any invertebrate group along the Florida coast has been done with 

crustaceans. While most of this research was done over 20 years ago, the combined results provide the 

bulk of what is known about the hardbottom community ecology along the east Florida coast. 

Decapods 

Camp et al. (1977) studied the nearshore ecology at Hutchinson Island, Florida (September 1971 to July 

1974). Sampling occurred during the day on five stations at depths of 7 to 11 m on sand and sand shell 

substrate. Three nearshore zones included 1) the shallow beach terrace, 2) crest of shallow off shore 

shoal, and 3) a deeper “trough” zone between them. More than 170 species of arthropods (of which 

113 were included in quantitative analysis) were recorded. Species recruitment curves never reached 

asymptote over the 2 years because of high species turnover rate and some poor grab efficiencies. 

Some 25 to 36 arthropods found only at the intertidal worm reef were unique to that habitat, with 

13 others also found offshore (Table 4.5). Highest densities of organisms were encountered from July to 

September, and lowest densities occurred January to March. The crabs Pachycheles monilifer

(porcellanid crab) and Pachygraphsus transverses (grapsid crab) were numerically dominant, but in terms 

of biomass, Menippe nodifrons (xanthid crab) was dominant. M. nodifrons was observed physically 

altering the reef by breaking off pieces, crushing them to expose the worms, and feeding on them. 

Shared species occurred infrequently or in low abundance offshore, indicating a distinct difference 

between worm reef and offshore faunas. Consequently, species associated with worm rock habitat were 

unique to that habitat compared to that offshore (Table 4.5). 

In a nearby study, Applied Biology, Inc. (1979) monitored a worm reef at the Florida Power & Light 

Company St. Lucie Power Plant from April 1976 to April 1979. Most of the sabellariid-associated fauna in 

this study were crustaceans, molluscs, and echinoderms. Species encountered, in order of abundance, 

were Menippe nodifrons, Pachycheles monilifer, Excorallana sexticornis, and Sphaeroma walki (isopods) 

and Pachygrapsus transverses (Tables 4.2, 4.6, and 4.7). The number of associated species collected 

was highest in July when water temperatures peaked and reef size was largest. Numbers and diversity of 

organisms was lowest in October to January, when water temperatures were decreasing and reef size 

was the smallest. 

Gore et al. (1978) investigated latitudinal changes in crustaceans in a 2-year study of four worm reef sites 

on the central east Florida coast from St. Lucie Inlet north to Sebastian Inlet (1974 to 1975). A total of 

96 species of decapod and stomatopod crustaceans representing 52 genera and 22 families was 

identified. Two surveys occurred, with sampling conducted both on the worm reef itself and in areas 

adjacent to the reef. Specifically associated with the worm reef were 24 common species, of which the 

first 11 species comprised 90% and the next 13 species comprised 4% to 7% of the collected samples. 

The remaining 3% (27 species) were considered uncommon (Table 4.8). The six most common species 

(80% of the samples) were the porcellanid crab Pachycheles monilifer, the grapsid crab Pachygraphsus 

transverses, the aphid pistol shrimp Synalphus fritzmuelleri, and the xanthid crabs Menippe nodifrons, 

Pilumnus dasypodus, and Panopeus bermudensis. P. monilifer and M. nodifrons are restricted to worm 

reefs along the east coast of Florida. The abundance of the dominant species was relatively similar at all 

sites, but the less dominant species varied among sites (Table 4.9). This variability is also indicated in 

that about 20 species appeared to be common to Gore et al. (1978) surveys (Table 4.5) and about 

31 species more occur in Gore et al. (1978) that are not listed in Camp et al. (1977). Furthermore, 

species richness was highest at St. Lucie Inlet south, intermediate (53%) towards the north from Walton 

Rocks to Fort Pierce Inlet, and lowest (13%) at Sebastian Inlet north. 
Section 2 4-20 

Organismal Assemblages of East Florida Shallow Hardbottom 

Table 4.5. Comparison of worm reef arthropods found in the vicinity of Hutchinson Island, Florida with 

arthropod species found offshore in the same area (From: Camp et al., 1977). ● denotes 

species presence. 

Stations 

I II III* IV V 

Depths (m) Species Found Only at Worm Reef Rank Species Found at Worm Reef 

and Offshore Sites 

8.4 11.2 7.1 10.9 10.9

Balanus tintinnabulum antillensis Tanystylum orbiculare ● ● 

Paranthura infundibulata Balanus trigonus 

(on Plagiusa depressa) ● ● ● ● ●

Sphaeroma walkeri Gonodactylus bredini ● ● ● ●

Bopyridae sp. D (on Petrolisthes armatus) Periclimenes americanus ● ● ● ●

Alpheus malleator Alpheus formosus ● 

A. nuttingi A. normanni ● ● ● ●

A. thomasi Synalpheus sp. B ● 

Synalpheus fritz muelleri Paguroidea sp. ● ● ● ●

Synalpheus sp. A Paguristes hummi ● ● ● ●

Calcinus tibicen Epialtus sp. ● ● 

Paguristes tortugae Mithrax forceps ● ● 

Pagurus brevidactylus Mithrax sp. ● ● 

Megalobrachium poeyi Pitho lherminieri ● ● 

Pachycheles monilifer 2 

Petrolisthes armatus 

P. galathinus 

Microphrys bicornutus 

Mithrax coryphe 

Eurypanopeus abbreviatus 

Menippe nodifrons 1 

Panopeus bermudensis 

Pilumnus dasypodus 

Xantho denticulatus 

Pachygrapsus transversus 2 

Plagusia depressa 

Yellow shading indicates species in common with Gore et al., 1978 study. 

* Station III had no hard substrate. 
Section 2 4-21 

Organismal Assemblages of East Florida Shallow Hardbottom 

Table 4.6. Abundance of associated decapods collected from sabellariid worm reef (Stations 9 and 10 

combined) at the St. Lucie Power Plant from April 1976 through April 1979 (From: Applied 

Biology, Inc., 1979). 

1976 1977 1978 1979 Species Apr Jul Oct Jan Apr Jul Oct Jan Apr Jul Oct Jan Apr 

Alpheus formosus 1 2 1 6 5 1 1 3 

Alpheus malleator 2 

Alpheus normanni 1 

Alpheus nuttingi 1 4 3 

Alpheus thomasi 2 2 

Alpheus sp. A 1 

Alpheus sp. 2 2 

Calcinus tibicen 1 

Epialtus sp. 1 

Eriphia gonagra 

Hippolytidae sp. 1 

Lysmata intermedia 2 1 

Lysmata wurdemanni 1 

Lysmata wurdemanni? 

Majidae sp. 1 

Menippe mercenaria 2 4 3 3 3 1 6 6 2 

Menippe nodi frons 26 11 172 27 19 114 198 159 19 58 99 93 35 

Microphrys bicornutus 1 

Pachycheles monolifer 107 101 42 3 66 71 39 86 74 36 1 15 69 

Pachygrapsus transversus 23 5 28 4 4 22 23 8 4 10 14 13 32 

Pagurus carolinensis 1 1 4 3 1 2 

Panopeus bermudensis 1 2 3 6 2 2 16 2 

Pelia mutica 1 

Petrolisthes armatus 1 1 1 1 

Petrolisthes galathinus 1 2 

Pilumnus dasypodus 2 19 11 14 10 22 3 3 3 

Pilumnus sp. 2 1 2 2 

Pitho lherminieri 2 

Pi tho sp. 1 

Plagusia depressa 1 1 

Synalpheus fritzmuelleri 6 6 2 2 3 74 15 4 13 1 1 2 

Synalpheus townsendi 1 

Synalpheus sp. 1 

Upogebia affinis 1 

Xantho denticulatus 2 1 1 

Xanthidae sp. 1 3 

Table 4.7. Comparison of associated decapod and stomatopod crustaceans from studies at Walton 

Rocks and the St. Lucie Power Plant (From: Applied Biology, Inc., 1979). 

 Applied Biology, Inc. 

St. Lucie Power Plant 

 

 

Smithsonian Institution 

Fort Pierce Bureaub

Walton Rocks 

 

Florida Department of 

Natural Resources 

Within 2 km South of 

St. Lucie Power Plant 

June 1975 1974 1975 

April 1976 

through 

January 1977 

April 1977 

through 

January 1978 

April 1978 

through 

January 1979 

Number of genera 22 15 11 12 18 14 

Number of species 30 20 13 15 29 18 

Diversity (d) --- 1.88 1.62 1.88 2.3 2.28 

Equitability (e) --- 0.1 0.31 0.33 0.24 0.39 

a

 Camp et al., 1977. 

b

 Gore et al., 1978 and R.H. Gore, pers. comm. 
Section 2 4-22 

Organismal Assemblages of East Florida Shallow Hardbottom 

Table 4.8. The 51 most recurrent species of decapod and stomatopod crustaceans encountered from 

sabellariid worm reefs in the Indian River region of Florida, 1974 to 1975 (From: Gore et al., 

1978). 

St. Lucie Walton Rocks Fort Pierce Sebastian 

Taxon 

1974 1975 1974 1975 1974 1975 1974* 

Abundance 60% to 80%

Spawning 

1. Pachycheles monilifer 127 163 256 None C MC Year round 

2. Menippe nodifrons 98 216 107 22 C MC May-Aug 

3. Synalpheus fritzmuelleri 54 64 42 None C MC Year round 

4. Pachygrapsus transversus 51 48 56 66 C MC Year round 

5. Pilumnus dasypodus 17 15 64 None C MC May-Aug 

6. Pitho lherminieri 14 4 None None C 

7. Mithrax coryphe 14 2 6 None C 

8. Gonodactylus bredini 11 4 None None C 

9. Pilumnus sayi 9 None 18 None C Jan-Aug 

10. Petrolisthes galathinus 8 1 14 None C Jan-Aug 

11. Alpheus formosus 8 None 8 None C Jan-Jul 

12. Pagurus carolinensis 8 1 1 None C May-Aug? 

13. Xantho denticulatus 7 3 None None C 

14. Gonodactylus oerstedii 7 None None None C 

15. Menippe mercenaria 6 20 24 3 C 

16. Synalpheus sp. A 6 None 4 None C 

17. Epialtus bituberculatus 6 None 10 None C Jan-Aug 

18. Alpheus nuttingi 5 4 9 None C Nov-Aug 

19. Microphrys bicornutus 4 None 4 None C 

20. Plagusia depressa 3 2 1 1 C May-Aug? 

21. Panopeus bermudensis 2 15 44 4 C MC May-Oct 

22. Alpheus thomasi 2 4 2 None C Jan-Aug? 

23. Synalphells minlls 2 None 7 None C 

24. Synalpheus townsendi 2 2 3 None C Jan-Aug 

25. Uhlias limbatus 2 None None None UC 

26. Processa fimbriata 2 None 1 None UC 

27. Metalpheus rostratipes 1 2 None None UC 

28. Lysmata intermedia 1 None 1 None UC 

29. Petrolisthes armatus 1 1 1 None UC 

30. Alpheus normanni 1 None None None UC 

31. Epialtus dilatatus 1 None None None UC 

32. Alpheus paracrinitus 1 None None None UC 

33. Megalobrachium soriatum 1 None None None UC 

34. Mithrax acuticornis 1 None None None UC 

35. Neopontonides beaufortensis 1 None None None UC 

36. Mithrax pleuracanthus 1 None None None UC 

37. Upogebia affinis 1 None None None UC 

38. Platyactaea setigera 1 None None None UC 

39. Micropanope granulimanus 1 None None None UC 

40. Mithrax hispidus None 2 None None UC 

41. Alpheus armillatus None 1 4 None UC 

42. Periclimenes americanus None 1 1 None UC 

43. Eurypanopeus dissimilis None 1 None None UC 

44. Lysmata wurdemanni None None 1 None UC 

45. Synalpheus brevicarpus None None 1 None UC 

46. Alpheus bouvieri None None 1 1 UC 

47. Paguristes tortugae None None 1 None UC 

48. Eurypanopeus abbreviatus None None 1 None UC 

49. Alpheus heterochaelis None None 1 None UC 

50. Panopeus occidentalis None None None 5 UC 

51. Panopeus herbstii None None None 1 UC 

* Sebastian Inlet station not sampled in 1975. C = common (97%); MC = most common (80%); UC = Uncommon (3%). 
Section 2 4-23 

Organismal Assemblages of East Florida Shallow Hardbottom 

Table 4.9. Species richness of decapod and stomatopod crustaceans collected from sabellariid worm 

reefs in the Indian River region of Florida, 1974 to 1975 (From: Gore et al., 1978). 

St. Lucie Walton Rocks Fort Pierce Sebastian* 

Most species rich 49% common with St. Lucie 56% common with St. Lucie 13% common with St. Lucie 

39 species 19 of 39 species 22 of 39 species 5 of 39 species 

* Sebastian Inlet station not sampled in 1975. 

Encompassing a larger geographical area along the Florida coast, van Montfrans (1981) studied decapod 

crustaceans associated with worm rock at five sites from Miami north to Cape Canaveral. There were a 

few species that accounted for the majority of individuals inhabiting intertidal and subtidal zones 

(Tables 4.10 and 4.11). The 4 most abundant of 15 species identified in the intertidal zone included 

Pachygrapsus transvenus, Menippe nodifrons, Panopeus bermudensis, and Pilumnus lacteus. In the 

subtidal zone, the 4 most abundant of 10 species included Synalpheus fritznuelleri, Pilumnus dasypodus, 

Menippe nodifrons, and Petrolisthes galathinus. Furthermore, species common to both subtidal and 

intertidal habitats included Menippe nodifrons, Panopeus bermudensis, Pachygrapsus transverses, 

Petrolisthes armatus, and Pilumnus lacteus. In terms of cross-shelf distribution, the diversity and 

abundance of decapods tended to increase with depth. In terms of latitudinal distribution in the subtidal 

zone, total numbers were highest at Deerfield Beach (610) but decreased towards the north at 

Hutchinson Island (394), Fort Pierce (202), and Satellite Beach (71). Moreover, total numbers decreased 

south of Deerfield Beach at Miami (296) (Table 4.12). Towards the southern two study sites (Miami and 

Deerfield Beach), Pilumnus dasypodus, Petrolisthes galathinus, and Synalpheus fritznuelleri were 

dominant, although their order of dominance varied. In the central two study sites (Hutchinson Island and 

Fort Pierce) the dominant species mix consisted of Menippe nodifrons (which occurs subtidally and 

intertidally), Synalpheus fritznuelleri, Pachycheles monilifer, and Petrolisthes galathinus. At the most 

northern study site (Satellite Beach), the species mix changed to the following three dominant species: 

Menippe nodifrons, Pachygrapsus transvenus, and Petrolisthes armatus. Many tropical species 

(Families: Alpheidae, Palaemonidae, Hippolytidae, Stenopodidae, Paguridae, Palinuridae, Porcellanidae, 

Xanthidae, Grapsidae, and Majidae) decrease in numbers and species from south to north, probably due 

to higher variability in sea surface temperature near Cape Canaveral versus Miami, and may be displaced 

further offshore in the north (van Montfrans, 1981). 

Table 4.10. Rank analysis of intertidal decapods from the east Florida coast (From: van Montfrans, 

1981).

Species Bio1. Index 

(3-pt. System) 

Frequency as 

One of Top 

Three Species in 

Nine Samples 

Frequency in 

Nine 

Samples 

Total Number 

Collected 

Greater 

Numbers 

Occurring in a 

Sample 

Overall 

Density/ 

1,000 m of 

Reef Sampled

Pachygrapsus transvenus 26 9 9 324 97 1.969 

Menippe nodifrons 11 5 5 113 91 0.687 

Panopeus bermudensis 4 3 4 23 17 0.140 

Pilumnus lacteus 4 2 2 11 10 0.067 

Alpheus bouvieri 2 1 3 7 3 0.043 

Clibinarius tricolor 2 1 2 3 2 0.018 

Eriphia gonagra 2 1 2 3 2 0.018 

Paguristes tortugae 2 1 1 2 2 0.012 

Pagurus carolinensis 2 1 1 20 20 0.121 

Petrolisthes armatus 2 2 4 10 6 0.061 

Alpheus malleator l 1 1 1 1 0.006 

Cyclograpsus integer 1 1 1 2 2 0.012 

Eurypanopeus abbreviatus 1 1 1 2 2 0.012 

Menippe mercenaria 1 1 4 5 2 0.030 

Plagiusa depressa 1 1 2 3 2 0.018 

Species highlighted in yellow occur in both intertidal and subtidal habitats. 
Section 2 4-24 

Organismal Assemblages of East Florida Shallow Hardbottom 

Table 4.11. Rank analysis of the dominant subtidal decapods from the east Florida coast (From: van 

Montfrans, 1981). 

Species Bio1. Index 

(3-pt. System) 

Frequency as 

One of Top 

Three Species 

in Nine 

Samples 

Frequency in 

Nine 

Samples 

Total Number 

Collected 

Greater 

Number 

Occurring in a 

Sample 

Overall 

Density/ 

1,000 m of 

Reef Sampled

Synalpheus fritznuelleri 11 6 7 443 131 1.727 

Pilumnus dasypodus 10 5 6 255 84 0.994 

Menippe nodifrons 9 4 7 330 241 1. 287 

Petrolisthes galathinus 6 3 4 399 308 1. 556 

Panopeus bermudensis 5 2 6 131 65 0.51 

Pachycheles monilifer 5 3 6 101 56 0.218 

Pachygrapsus transversus 5 2 5 34 15 0.132 

Petrolisthes armatus 1 1 7 27 6 0.105 

Pilumnus lacteus 1 1 3 87 62 0.339 

Lysmata wurdemianni 1 1 2 11 7 0.027 

Species highlighted in yellow occur in both intertidal and subtidal habitats. 

Table 4.12. Total number of the three most common subtidal decapod species at stations along the east 

Florida coast (Adapted from: van Montfrans, 1981).

Station A 

Dade 

Miami 

Station B 

Broward 

Deerfield Beach 

Station E 

St. Lucie 

Hutchinson Island

Station C 

St. Lucie 

Fort Pierce 

Station D 

Brevard 

Satellite Beach 

Pilumnus 

dasypodus 106 Petrolisthes galathinus 328 Menippe nodifrons 247 Pachycheles 

monilifer 71 Menippe 

nodifrons 44 

Petrolisthes 

galathinus 103 Synalpheus fritznuelleri 165 Synalpheus fritznuelleri 137 Petrolisthes galathinus 71 Pachygrapsus 

transvenus 20 

Synalpheus 

fritznuelleri 87 Pilumnus dasypodus 117 Pachycheles monilifer 10 Synalpheus fritznuelleri 60 Petrolisthes 

armatus 7 

Totals 296 610 394 202 71 

All common to the 

subtidal zone 

All common to the 

subtidal zone 

M. nodifrons common to

both zones 

All common to the 

subtidal zone 

All common to both 

subtidal and intertidal 

zones 

Southern sites Middle sites Northern range 

Species highlighted in yellow occur in both intertidal and subtidal habitats. 

Reed et al. (1982) studied the composition of decapod crustaceans and the total number of individuals 

per species on east Florida oculinid coral reefs. A total of 50 species was identified with four locations of 

varying depth being examined: Fort Pierce Inlet (6 m), north of Fort Pierce (27 m), south of Fort Pierce 

(42 m), and offshore east of Fort Pierce Inlet (80 m depth). Species showed strong clustering by station 

with a gradual decrease in density as depth increased (Table 4.13). At the 6-m depth, there were 

discrete species clusters that were considerably different from all the other stations. Stations at 42- and 

80-m depths showed little species differences. Species composition was relatively stable at each depth, 

and there were no dramatic seasonal overturns of assemblages. However, there was a degree of trophic 

portioning between the shallow station (Fort Pierce: 6 m), which was numerically dominated by the 

filter-feeding porcellanid crab Megalobrachium soriatum, and the 27-m station, which was dominated by 

the carnivorous Pagurus carolinensis. Four species that were found at all study sites included 

Megalobrachium soriatum and Synalpheus townsendi (Tables 4.13 and 4.14). 
Section 2 4-25 

Organismal Assemblages of East Florida Shallow Hardbottom 

Table 4.13. Numerical abundance of decapod species at Oculina varicosa coral reef stations at depths of 

6, 27, 42, and 80 m (Adapted from: Reed et al., 1982). 

Number 

Abundance 

6 m 

Fort Pierce Inlet 

27 m 

North of Fort Pierce 

42 m 

South of Fort Pierce 

St. Lucie Inlet 

80 m 

Offshore East of 

Fort Pierce 

1 Megalobrachium soriatum Pagurus carolinensis Pagurus carolinensis Pagurus carolinensis 

2 Pachycles monilifer Megalobrachium soriatum Megalobrachium soriatum Megalobrachium soriatum 

3 Mithrax forceps Synalpheus townsendi Pagurus piercei Pagurus piercei 

4 Thor manningi Mithrax forceps Synalpheus townsendi Synalpheus townsendi 

5 Synalpheus fritzmuelleri Pelia mutica Periclimenes iridescens Galathea rostrata 

6 Synalpheus cf. townsendi Stenorhynchus seticornis Micropanope scuptipes 

7 Micropanope nuttingi Thor manningi 

8 Paguristes tortugae Pseudomedaeus distinctus 

9 Pseudomedaeus agassizii 

10 Nematopaguroides pusillus 

11 Euchirograpsus americanus 

Table 4.14. Percent occurrence of decapod species at Oculina varicosa coral reef stations at depths of 

6, 27, 42, and 80 m (Adapted from: Reed et al., 1982). 

Percent 

Occurrence 

6 m 

Fort Pierce Inlet 

27 m 

North of Fort Pierce 

42 m 

South of Fort Pierce 

St. Lucie Inlet 

80 m 

Offshore East of 

Fort Pierce 

Pagurus piercei 

~100 Synalpheus townsendi Pagurus carolinensis 

Megalobrachium soriatum Pagurus carolinensis 

Galathea rostrata 

Megalobrachium soriatum Pagurus carolinensis ~90 Mithrax forceps Pagurus piercei Micropanope scuptipes 

~80 Pachycles monilifer Synalpheus townsendi 

Thor manningi 

~70 Synalpheus townsendi Periclimenes iridescens Pseudomedaeus distinctus 

Nematopaguroides pusillus 

Thor manningi Megalobrachium soriatum ~60 Synalpheus fritzmuelleri Megalobrachium soriatum Euchirograpsus americanus 

Mithrax forceps 

Pelia mutica 

Stenorhynchus seticornis 

Micropanope nuttingi 

~50 Synalpheus cf. townsendi

Paguristes tortugae 

Pseudomedaeus agassizii 

≤40 Rest Rest Rest 

Crustacean species found on 6-m oculinid corals include Epialtus spp., Pagurus spp., and Synalpheus

spp. (which is often associated with sponges and algae) (Reed et al., 1982). Also, there were 14 species 

that appeared to be shared between oculinid and sabellariid worm reefs mentioned in Gore et al. (1978). 

Furthermore, Reed et al. (1982) reported that on oculinid reefs, Pachycheles monolifer and Synalpheus 

fritzmuelleri were abundant, Pagurus carolinensis was common, and that Syalpheus minus, S. townsendi, 

and Megalobrachium soraitum were rare on sabellariid reefs. Species common on the sabellariid reefs 

but rare on oculinid reefs included Pilumnus dasypodus, Petrolisthes galathinus, Epialtus sp., Alpheus 

formosus, and Synalpheus cf. fritzmulleri. 

The gradient in species assemblages was attributed to a gradient of environmental factors across the 

shelf (Table 4.15). At a depth of 6 m, the environment is characterized as having variable wave surge 

and sedimentation. The temperature was about 25°C and varied less relative to other sites, which 

accounted in part for the highest diversity of species – 11 endemic and uniquely associated with this 

depth. At 42 m, temperature was more variable, averaging about 18°C, with relatively less wave energy 

and sedimentation. As a result, diversity and number of endemic species was relatively low but individual 

numbers were highest. At the 80-m depth, the physical parameters were similar to those at the 42-m 

station, but there were relatively high numbers of species and endemic species but lowest individual 
Section 2 4-26 

Organismal Assemblages of East Florida Shallow Hardbottom 

numbers of each species. Temperature changes caused by upwelling events that occur periodically were 

considered to be an important factor controlling the community structure on these oculinid reefs. 

Table 4.15. Physical parameters measured between 1976 and 1979 at Oculina varicosa coral reef 

stations at depths of 6, 27, 42, and 80 m. Range (mean and ± 1 standard deviation) 

(From: Reed et al., 1982). 

Parameter 6 m 

Fort Pierce Inlet 

27 m 

North of Fort Pierce 

42 m 

South of Fort Pierce 

St. Lucie Inlet 

80 m 

Offshore East of 

Fort Pierce 

Salinity (%) 28.2-36.4 (32.5 ± 3.4) 35.9-36.2 (only 2 values) 35.7-36.4 (36.0 ± 0.2) 

Temperature (°C) 13.7-31.0 (24.6 ± 3.1) 8.0-27.8 (18.4 ± 2.7) 7.4-26.7 (16.2 ± 3.7) 

Current (cm/sec) 0-70.5 (8.9 ± 2.1) 0-49.5 (8.7 ± 2.4) 0-58.5 (8.6 ± 1.7) 

Visibility (m) 0-9 (3 ± 2) 0-21 (9 ± 5) 0-25 (9 ± 5) 

Sedimentation 

(mg/cm2

/day) 137-1,640 (846 ± 593) 

No data available 

No data available 15-78 (53 ± 34) 

Further south along the coast, Cummings (1994) reported the presence of six species of crustaceans 

associated with the colonization of a newly constructed artificial reef structure in the nearshore 

(~8 m depth) area off Boca Raton, Florida (April 1988 to July 1990). Decapod species observed from 

December to January 1988 included Eurypanopeus abbreviatus and Percnon gibbesi. From July to 

August 1989, Mithrax forceps, Pagurus carolinensis, and Percnon gibbesi were identified. In addition to 

these species, Sultzman (1990, 1997) encountered penaeid shrimp and blue crabs in the nearshore 

areas along the Florida coast. 

Isopods and Amphipods 

Nelson and Demetriades (1992) sampled worm rock from rock jetties at Sebastian Inlet, Florida, for a 

period of 1 year (April 1984 to March 1985) at daytime low tides. They collected six isopod species, of 

which Paradella dianae (53,000/m2

) and Sphaeroma walkeri (35,000/m2) represented 98% of the total 

number of 6,136 isopods present (Table 4.16). Two peaks in abundance occurred for both species 

during the year (February-March and August-October). Recruitment was year-round (both species), and 

growth was bimodal for P. dianae (mode 1.1 to 1.5 mm size class in all months; 2.7 to 3.1 mm some 

months; shifted smaller [2.3 to 2.7 mm] or larger [3.1 to 3.5 mm]) and bimodal and skewed unimodal for 

Sphaeroma walkeri (some months mode 0.85 to 1.75 mm size class; shifted smaller [1.75 to 2.65 mm] or 

larger [2.65 to 3.55 mm] some months). Total monthly isopod density was negatively correlated with 

percent algal coverage on worm reef, and S. walkeri density was negatively correlated with air and water 

temperature on the north jetty site (which received more sunlight) but not on the south jetty site. Pardella 

danae density was not significantly correlated with physical parameters. Filamentous algal coverage by 

species such as Enteromorpha sp. tended to restrict access to suitable crevices in the worm rock. Nelson 

and Demetriades (1992) also collected 10 amphipod species, of which Hyale perieri (22,000/m2

), 

Elasmopus pectinicrus (10,000/m2

), and Ampithoe pollex (4,000/m2) represented 91% of the total number 

of 3,801 amphipods (Table 4.16). Maximum abundance was observed to occur in late summer and early 

spring for Hyale perieri and was negatively correlated with salinity. For E. pectinicrus, a single maximum 

occurred during October to November, and abundance for A. pollex appeared to be lowest during 

November to January and was positively correlated with air temperature. 

S. walkeri is more widely distributed than P. danae along the east Florida coast (Camp et al., 1977). 

Carlton and Iverson (1981) described its natural history and indicate that distribution is 

temperature-limited at a minimum surface winter isotherm of 15°C, which is typical at Sebastian Inlet 

(December to January). P. danae is typically cryptic in its habitat and found under stones, in empty 

barnacle tests, under chitons, in empty worm rock tubes, or in crevices of worm rock. 
Section 2 4-27 

Organismal Assemblages of East Florida Shallow Hardbottom 

Table 4.16. Abundance of isopods and amphipods from sabellariid worm rock with percentage of 

contribution of species to total abundance of each order at Sebastian Inlet, Florida, 

April 1984-March 1985 (From: Nelson and Demetriades, 1992). 

Species Abundance Percentage of 

Relative Abundance 

Isopoda 

Paradella dianae 4,244 69.0 

Sphaeroma walkeri 1,757 28.7 

Paranthura infundibulata 79 1.3 

Jaeropsis sp. nov.1 39 0.7 

Excorallana spp.2 19 0.3 

Amphipoda 

Hyale perieri 2,189 57.5 

Elasmopus pectinicrus 966 25.4 

Ampithoe pollex 396 10.4 

Corophium acutum 145 3.8 

Ericthonius sp. 55 1.4 

Jassa cf. falcata 21 0.5 

Ampithoe longimana 18 0.5 

Stenothoe georgiana 11 0.3 

Podocerus brasiliensis 2 0.1 

Cymadusa compta 1 0.1 

1

 Undescribed species, T.E. Bowman, personal communication. 

2 Includes two species, one of which is probably Excorallana sexlicornis. Positive identifications could not be made 

because few fully differentiated males were present (T.E. Bowman, personal communication). 

4.3.1.3 Reproduction and Life History 

Most crustaceans have separate sexes and reproduce sexually. Depending on the species, fertilization 

can be internal or external, with eggs being either shed directly into the sea or brooded by the females 

(most common with crabs and shrimp). In either case, most crustaceans produce larvae (nauplius or 

zoea) that eventually end up as part of the plankton. They can be lecithotrophic or planktotrophic and can 

be in the plankton from a few weeks (i.e., xanthid crabs, penaeid shrimp) to almost a year (i.e., spiny 

lobster) (Lyons et al., 1981; Young and Chia, 1987; Florida Wildlife Research Institute [FWRI], 2007a). In 

contrast, isopods, amphipods, and other pericarids lack larval stages. Their young emerge as miniature 

versions of the adults. Consequently, there is a wide range of dispersal capabilities among crustaceans. 

Upon recruitment, some become reproductive quickly and are short-lived like the amphipods and isopods 

(Nelson and Demetriades, 1992), while others, such as stone crabs (Lindberg and Marshal, 1984; Bert, 

1992; FWRI, 2007b) and spiny lobster (Lyons et al., 1981), may live several years. 

General reproduction and life history are fairly well known for such groups as penaeid shrimp and blue 

crabs. Penaeid shrimp have a short life span of about 1.5 years and spawn offshore in deeper waters 

(FWRI, 2006). The larvae develop in the plankton and move back into the salt marshes and tidal creeks. 

Blue crabs (Callinectes sapidus) have a life span of about 4 years, reaching sexual maturity in 1 year 

(Tagatz, 1968a). Females mate once in their lifetime during March to July or October to December 

(Tagatz, 1968a,b). After mating in the upper reaches of estuaries, females migrate to the mouth of the 

estuary and nearshore areas to spawn (FWRI, 2007c). Zoea drift in continental shelf waters for as long 

as 30 to 45 days (FWRI, 2007c), but this can vary by species and with temperature. 

The Cuban stone crab (Menippe nodifrons) spawns from May to August (Gore et al., 1978), and the 

megalope stage is attainable within 16 to 17 days at 30°C in the laboratory, versus 28 to 37 days at 20°C 

(Scotto, 1979). Cuban stone crabs are closely allied to their commercial exploited counterpart Menippe 

mercenaria, which also has abbreviated zoeal stages (five or atypically six), depending on the ambient 
Section 2 4-28 

Organismal Assemblages of East Florida Shallow Hardbottom 

water temperature (Scotto, 1979). Ong and Costlow (1970) suggested 30°C was the optimum survival 

temperature for M. mercenaria larvae, with optimum salinity of 30 to 35 ppt (megalopal stage was 

reached in 14 days, first crab in 21 days, with 60% to 72% larval survival rate). Highest survival rate with 

M. nodifrons (15%) occurred at 20°C (megalopal stage was reached in 28 to 34 days, first crab in 45 to 

49 days, but this can be prolonged; megalopal stage reached in ~37 days, first crab in ~52 days, with the 

atypical sixth zoeal stage). Stone crabs may live as long as 7 years (FWRI, 2006). Andryszak and Gore 

(1981) reported in their study of larval development of the deep water xanthid Micropanope sculptipes

that it had four larval stages, like the majority of xanthid species. As a result, species with fewer larval 

stages would indicate that they spend less time as part of the plankton. 

Spiny lobsters (Panulirus argus) spawn in late spring and early summer on the outer reefs and offshore 

(Table 4.17). The larvae (phyllosoma) live offshore for 6 to 12 months and are solitary, observing a 

diurnal vertical migration pattern (occurring shallower at night and deeper during the day). Post-larval 

(puerulus) recruitment occurs year-round, and the larvae move from the open ocean towards nearshore, 

moving in the surface 1 m of water. Movement is aided by wind-driven surface currents and/or flood tides 

at night following the new moon. When they are >15 to 20 mm carapace length (CL), they move into 

crevices and nocturnally feed on invertebrates, mainly molluscs (depth <5 m). As growth increases to 

20 to 50 mm CL (7 to 10 months), they move to deeper water (depths of 10 to 20 m), 50 to 80 mm CL 

(1 year) (depths of 100 to 500 m), and >80 mm CL (15+ years) (depths of 500 to 1,000 m). Typically, 

1-year-old subadults are found in 3- to 10-m water depths. Most crustacean larvae are subsequently 

returned to estuarine and nearshore areas by tides and wind-driven currents, which may cause some 

unexpected upwelling events (Pitts and Smith, 1997; Pitts, 1999). 

Along the east Florida coast, some crustaceans spawn year-round, while others do so only seasonally 

(Tables 4.8 and 4.17). Notable year-round spawners are Pachycheles monifer, P. transversus, and 

Synalpheus fritzmuelleri. Ovigerous females of P. dianae and S. walkerii reported by Nelson and 

Demetriades (1992) were present year-round, so they probably spawn year-round as well. In contrast, 

Menippe nodifrons, Pilumnus spp., Pagurus sp., Petrolisthes galathinus, and Alpheus spp. appear to 

spawn seasonally during the spring or summer time period. 

Reed et al. (1982) suggested that recruitment of Oculina coral reefs in shallow areas (6 m depth) along 

the east Florida coast may occur from two directions, the surrounding algae-sponge habitat and adjacent 

sabellariid worm reefs. The authors provided examples for species colonization that may occur from 

inshore to offshore, and vice versa: 1) Pachycheles monilifer, which was dominant on sabellariid worm 

reefs, was also restricted to 6-m depth on oculinid reefs; 2) Megalobrachium soriatum, which was the 

second most common species on oculinid reefs and rare on worm reef according to Gore et al. (1978), 

was found at all oculinid stations (6-, 27-, 42-, and 80-m depths); 3) the following species were observed 

on both sabellariid worm reef and 6-m oculinid reef but not at the deeper stations: Synalpheus 

fritzmuelleri, Pilumnus dasypodus, Petrolisthes galathinus, Alpheus formosus, and Epialtus sp.; and 

finally, but considered least likely, 4) colonization could occur as the larvae settle out of the plankton from 

offshore at 80-m depths moving shoreward, and the perseverance of various species would be controlled 

by selective elimination as a result of periodic cold water upwelling events. Recruitment of dominant 

isopods associated with worm rock occurred in most months (Nelson and Demetriades, 1992). 
Section 2 4-29 

Organismal Assemblages of East Florida Shallow Hardbottom Table 4.17. Summary of the life history characteristics of the ontogenetic stages of spiny lobster (Panulirus argus) (Adapte

http://bio.fsu.edu/~herrnlab/Life_History_Table.pdf). Life Stage Phyllosoma Puerulus Algal phase Post-algal Juvenile 

Size 1 to 10 mm 5 to 6 mm CL 6 to 20 mm CL 20 to 50 mm CL 50 to 80 mm CL Duration 6 to 9 months 1 to 2 weeks 2 to 3 months 7 to 10 months 1 year Color Transparent body Transparent body Cryptic: banded disruptive Semi-cryptic: spotted Semi-cryptic: spoHabitat Open ocean Offshore moving inshore Macroalgae, seagrass Bays, hardbottom sponges, holes Bays, hardbottom patch reefs, amid seagrass holes Food Soft-bodied zooplankton Non-feeding stage Generalist: invertebrates (especially molluscs) Same Same Forage grounds Upper 200 m Non-feeding stage Macroalgal patches Grass beds, rocks Grass beds, rockHome range Unknown Not available <5 m 10 to 20 m 100 to 500 m Predators Pelagic fishes Coastal fishes, jellyfishes Small fishes, crabs, octopus Fishes, crabs, octopus Octopus, large fis

stingrays, man 

Antipredation strategy Sparseness Fast-moving, crypsis Camoflage, shelter use Shelter use, aggregation Shelter use, aggregation, cooperative defeActivity Diurnal vertical migration Nocturnal Nocturnal only Nocturnal only Nocturnal crepus

(diurnal at mass migration) 

Sociality Solitary Solitary Solitary, asocial Gregarious (forage as individuals) Gregarious (forag

individuals) 

CL = Carapace length. 
Section 2 4-30 

Organismal Assemblages of East Florida Shallow Hardbottom 

4.3.2 Gastropods and Other Molluscs 

4.3.2.1 Diversity and Trophic Function 

There is a high diversity and abundance of gastropods on hardbottoms along the east Florida coast. 

Reed and Mikkelsen (1987) studied the molluscan community associated with the scleractinian coral 

Oculina varicosa at four sites of varying depth. They identified 230 species-level taxa, including 

155 gastropods, 68 bivalves, 1 scaphopod, 5 polyplacophorans, and 1 cephalopod (Table 4.4 and 

Appendix B, Table B.2). The observed taxa utilized four modes of existence: motile species (47%), 

symbiotic (32%), epilithic (18%), and endolithic (3%). There may be as many as 40 gastropod species 

that are rare and endemic to small nearshore reaches of Florida coastline (USFWS, 1999), specifically 

beach-rock outcrops along Palm Beach and Martin Counties (Petuch, 1988). 

Gastropods are likely very important in food webs along the east Florida coast. They are important as 

predators of various invertebrates and small fishes (McGraw and Gunter, 1972, Carriker, 1978; Reed and 

Mikkelsen, 1987; Watanabe and Young, 2006). Watanabe and Young (2006) indicated that the main 

predators of P. lapidosa were the dog whelk Stramonita haemastoma and decapod crustaceans (Gore et 

al., 1978), although they also observed low numbers of the non-shell-boring gastropods Pollina tinca

(Buccinidae) and Leucozonia nassa (Fasciolaridae) feeding on P. lapidosa. Stramonita haemostoma

traditionally feeds on oysters, however, at Walton Rocks had developed a much longer proboscis to feed 

on worms than conspecifics that fed on bivalves (Watanabe, 2002). Boring gastropods feed by scraping 

the surface of shells and also by applying secretions that dissolve the shell, or by injecting toxins 

(McGraw and Gunter, 1972; Carriker, 1978). Additionally, gastropods can be a major prey item for other 

invertebrates such as the stone crab Menippe mercenaria occasionally observed on worm reefs, portunid 

crabs (Butler, 1985; Richardson and Brown, 1992), spiny lobsters (Panulirus argus) (Sultzman, 1990), 

blue crabs (FWRI, 2007c), and other snails such as Pisania tincta (Buccinidae) (Watanabe, 2002), as well 

as fishes (puffers, sheepshead, black sea bass, porgies, yellowtail snapper) (South Carolina Marine 

Resources Research Institute and Duke University Marine Laboratory, 1982; Cummings, 1994; USFWS, 

1999). 

4.3.2.2 Latitudinal and Cross-shelf Distribution 

There is not much information on how gastropod diversity and abundance change with latitude and depth 

along the east Florida coast. Petuch (1987) reported that Cerithium lindae, Nerita lindae, and Modulus 

papei and two unnamed periwinkle species were associated with these nearshore habitats. Furthermore, 

it is possible to find Thias rustica, T. deltoidea, and T. floridana (also Purpura patula) along the east 

Florida coast. Two species of prickly winkle (Tectarius antonii and T. nodulosus), as well as the abundant 

Cenchrus muricatus, occur along Florida. Aplysia increases in abundance and number of species moving 

south towards the Keys. Nerites and Siphonarias are tropical, and few are encountered that survive the 

cold winter months. Petuch (1987) mentions endemic intertidal species, including several new species of 

small gastropods, some of which include Nerita lindae and Cerithium lindae from Martin, Palm Beach, 

and Broward Counties and some yet unnamed findings. In the rocky intertidal zone along Palm Beach 

County, it is possible to find Nerita peleronta, N. versicolor, N. tesselata, N. fulgurans, and N. lindae

(E. Petuch, Florida Atlantic University, pers. comm.). 

Applied Biology, Inc. (1979) monitored a sabellariid worm reef at the St. Lucie Power Plant (April 1976 to 

April 1979) and found 17 gastropod and 5 bivalve species associated with the reef. The four most 

common species of gastropods included Anachis avara, Mitrella argus, Anachis floridana, and Barleeia 

tincta (Table 4.2). As with other fauna observed, the number of gastropod species collected was highest 

in July when water temperatures peaked and reef size was largest (July 1977). Numbers and diversity of 

organisms were lowest from October to January when water temperatures were decreasing and reef size 

was the smallest. McCarthy (unpublished) noted the presence of Stramonita haemostoma (dog whelk) 

and Aplysia dactylomela (spotted sea hare) in observations at Bathtub Reef (2002 to 2003). 

S. haemostoma was significantly abundant all over the worm reef, but only during fall and early winter. 
Section 2 4-31 

Organismal Assemblages of East Florida Shallow Hardbottom 

Reed and Mikkelsen (1987) conducted studies of molluscs on the east Florida coast associated with 

Oculina variosa coral reef at four sites and depths (inner shelf 6 m, mid-shelf 27 m, outer shelf 42 m, and 

shelf edge 80 m). At the 6-m depth, Parviturboides interruptus was the most abundant species, 

representing 54% of specimens. In comparison, all other stations were dominated by at least five species 

that made up 50% of the specimens observed. Chione grus was dominant at 27 m (13%), Lithophaga 

bisculata was dominant at 42 m (16%), and Costoanachis lafresnayi was dominant at 80 m (24%) 

(Figure 4.18). 

Figure 4.18. Species composition at each of four Oculina stations (From: Reed and Mikkelsen, 1987). 

Numbers in parentheses are percent of total individuals for each station. 

In addition, three distinct species groupings were determined. One species group common to the 6-m 

depth was dominated by Parviturboides interruptus, Diplothyra smithii, Rissoina bouryi, and two species 

of Odostomia. The second, at 80-m depth, was dominated by Metaxia rugulosa, Calliostoma roseolum, 

and two other species of Odostomia. Finally, a third species grouping consisting of species common 

throughout the shelf was dominated by Lithophaga bisculata, Calliostoma sparsa, and Fargoa bushiana. 

The density of organisms at 6 m was significantly similar to that at the 80 m depth station but was 

significantly higher than that encountered at the 42 m depth (Table 4.4). Seasonal variations in 

community composition were minimal at 80 m because of high similarity of within-station fauna and low 
Section 2 4-32 

Organismal Assemblages of East Florida Shallow Hardbottom 

fluctuations in ordination scores. Also, at depths greater than 6 m, seasonal fluctuations in numbers of 

individuals and species were not as pronounced. At 6-m sites, these parameters varied according to 

temperature cycles, with a peak in numbers and species in June. This was due primarily to high densities 

of Parviturboides interruptus, Chione grus, and Diplothyra smithii and influx of rare species, respectively. 

Cold water upwelling and cooler average temperatures may have accounted for more eurythermic 

tropical, temperate, and boreal species at the shelf edge (80 m) than on the inner shelf (6 m). Regarding 

trophic structure, species assemblages at 80 m were distinct from those at 6 m, in that specimens 

collected from the 6-m depth were predominantly herbivores and detritovores. Those collected from 

80-m depths were predominantly carnivorous species. Habitat structure at the 80-m depth consisted of 

more dead and living coral than the other sites and, as a result, provided more space for epilithic molluscs 

and epizooiic (sponges, tunicates, and hydroids) and carnivorous species to colonize. At the 6-m depth, 

more boring endolithic species of molluscs prevailed, with densities five times that compared to the 42- or 

80-m stations (Appendix B, Table B.3). 

4.3.2.3 Reproduction and Life History 

Gastropod species sexually reproduce but can be dioecious or hermaphroditic. Fertilization can be 

internal or external, producing offspring that develop directly or indirectly. In direct development, offspring 

often are encapsulated in egg cases that are attached to the seafloor. Juveniles hatch out of the cases 

as miniature adults. In indirect development, a trochophore (only archaeogastropods such as limpets, 

nerites, and turbans) or veliger larva (most other gastropods) is produced. In the latter case, veligers can 

be lecithotrophic and short-lived (days to weeks) in the plankton or planktotrophic and long-lived (up to 

3 months) in the plankton. 

Little is known about seasonal spawning and recruitment patterns of gastropods along the Florida coast. 

These larvae are among the most abundant nearshore zooplankton, but their occurrence is likely to be 

both local and brief, and information about larval ecology is limited (Reed and Mikkelsen, 1987; Johnson 

and Allen, 2005). However, community persistence seems to be influenced strongly by physical factors 

such as the stability and sustainability of adequate substrate, reduced wave energy, and/or severe storm 

surf (Woodley et al., 1981; Sousa, 1984), as well as adequate food supply. Also, habitat that is not prone 

to severe desiccation like intertidal habitats (as a result of low tides and high temperature maxima) can 

support more snails (Newell, 1979). In addition, Watanabe (2002) noted that low abundance of 

encrusting flora and fauna, presence of filamentous algae, and smooth worm reef mounds (dead worm 

reef) tended to increase stress and mortality to intertidal invertebrates, leading to a reduction in food 

supply for the snails. The physical environment is markedly different between inner and outer shelf reefs. 

At the 6-m depth, there were more waves, sedimentation, light, and constant temperature conditions that 

favored algal growth and herbivores. At a depth of 80 m, temperature fluctuations are as much as 

13°C in 48 hours, and Reed and Mikkelsen (1987) speculated that recruitment of molluscan larvae from 

species pools outside of the Oculina reefs may be greater at the shelf edge reefs than at the inner shelf 

because they are closer to the warm Florida Current, which supplies tropical species of larvae from the 

south year-round. Moreover, outer shelf reefs are close to the deep, cool counter-current from the north 

that flows through the Florida Straits. The latter supplies temperate and boreal species to the shelf edge 

reefs. Larval distribution is probably greatly affected by numerous upwelling events caused by wind 

stress and tides (Pitts and Smith, 1997; Pitts, 1999; McCarthy, 2005). 

Watanabe (2002) conducted studies on the dog whelk Stramonita haemostoma occurring at Marineland 

(rock boulders) and Bathtub Reef (worm rock) along the east Florida coast (1999 to 2001). Usually, 

spawning occurs from April to August (Butler, 1985). However, this can be affected by diet. Watanabe 

(2002) found that snails feeding on their normal diet of oysters reproduced earlier (May to August) 

compared to those feeding on P. lapidosa (July to September). Data from Marineland indicated that 

S. haemostoma is capable of producing from 226 to 270 egg capsules per snail, and as many as one to 

six clutches over the summer, with some individuals spawning multiple times. Egg capsules contain 

larvae that hatch into veligers after 10 to 27 days and then become part of the plankton for several 

months (Scheltema, 1971; Dobberteen and Pechenik, 1987). Each egg capsule can contain from 836 to 

1,766 larvae (Watanabe, 2002). In general, the life span of these gastropods is about 5 years, with 
Section 2 4-33 

Organismal Assemblages of East Florida Shallow Hardbottom 

seasonally-induced settlement throughout their range (Gunter, 1979; Butler, 1985). However, the 

production of snails was lower on the Bathtub Reef worm reef (life span of 1 year) compared to rock 

boulders at Marineland (significantly longer). The community of snails on worm rock consisted of a new 

single cohort generated each year (recruited September to October) that completely died off by July to 

August, after 9 to 10 months. In addition, P. lapidosa recruitment and demise occurred in the same time 

frame (September to October and July to August, respectively) (McCarthy, 2001; Watanabe, 2002). 

No reproduction occurred on the worm rock at Bathtub Reef; however, at Marineland, the snail 

community was more persistent, with multiple cohorts (new recruits arriving from October to November) 

and reproduction and survival through fall and summer. Watanabe (2002) postulated that this may be 

due to availability of food throughout the year and the ability to survive harsh summer heat and weather 

conditions. Consequently, this was reflected in snail density, which was lower at Bathtub Reef 

(0.6 to 5.5 individuals/m2

/yr), representing a single cohort each year, compared to other locals with 

multiple cohorts per year, i.e., Marineland (16.1 ± 19.8 individuals/m2

/yr), Louisiana oyster beds 

(2.9 to 37.9 individuals/m2

/yr) (Brown and Richardson, 1987), and Gulf (2.7 to 27 individuals/m2/yr) 

(Butler, 1985). 

In southeast Florida, Cummings (1994) studied the recruitment and community development associated 

with a newly constructed artificial reef (April 1988 to July 1990) off Boca Raton, Florida. A total of 

64 invertebrate species was recorded during the study, of which three were gastropod species and four 

were bivalve species (Table 4.3). Invertebrate species diversity was significantly higher in July 1990 than 

in December 1988 to January 1989. However, the seasonality of invertebrates was not confirmed 

because of a temporary and unexpected burial of four of six sampling locations. Generally, the 

invertebrate community never seemed to reach an equilibrium and stayed in an early stage of 

development because of frequent physical disturbance. 

4.3.3 Echinoderms 

4.3.3.1 Diversity and Trophic Function 

Echinoderms can be abundant along some east Florida coast NHBs. While echinoids are often most 

visible out on reefs, there are a number of holothurians, ophiuroids, asteroids, and crinoids that can be 

encountered along the coast (Appendix B, Table B.1). Overall, most echinoderms do not provide 

significant shelter for higher numbers of species. Some exceptions to this may occur with small animals 

such as shrimp, mysids, copepods, porcellanid crabs, brittle stars, blennies, gobies, and pearlfish 

(Randall et al., 1964: Schoppe, 1991). However, some echinoderm species may be very important 

herbivores that influence abundance or diversity of macroalgae that in turn affects the amount of either 

1) shelter or food for other species, and/or 2) space availability for recruitment of other sessile organisms 

such as stony corals (see Duffy and Hay, 2001 for a review). In the more northern sections of east 

Florida NHB, the echinoid Echinometra lucunter is very abundant and significantly reduces biomass of 

macroalgae and cyanobacteria (D. McCarthy, Jacksonville University, Jacksonville, FL, pers. observ.). 

Such removal of macroalgae and cyanobacteria may be important in providing recruitment opportunities 

for sessile invertebrate such as Oculina and Siderastrea. Recent observations of removal of macroalgae 

by urchins were documented during monitoring of the Indian River County Ambersand mitigation reef. 

Observations indicated an 8-fold increase in urchin populations on the mitigation artificial reef between 

2006 and 2008 where removal of macroalgae and mud cover increased recruitment opportunities for the 

scleractinian Oculina (Miller and Kosmynin, 2008; Coastal Eco-Group Inc., 2009). Other echinoderms 

such as some holothurians and brittle stars are deposit-feeders that may convert detrital food sources into 

macroorgansim biomass that is available to higher trophic levels and enhance sediment resuspension 

and nutrient exchange with the water column (Bertness, 2007). 

Echinoderms can be important as a food sources to key species in a number of marine ecosystem food 

webs (see Duffy and Hay, 2001 for a review) but have not been studied in this sense along the east 

Florida coast. It is very likely that sea urchins are important prey components of food webs as there are 

several predators found along the coast that are relatively abundant and known to prey on sea urchins. 

Examples of these predators include spiny lobsters, spider crabs, grunts, jacks, wrasses, triggerfishes, 

and birds (Abbott et al., 1974; Hendler, 1977). 
Section 2 4-34 

Organismal Assemblages of East Florida Shallow Hardbottom 

4.3.3.2 Latitudinal and Cross-shelf Distribution 

How abundance and diversity of echinoderms change along east Florida coast NHBs is unclear for most 

of this group. The clearest patterns occur with echinoids, with a much higher biomass of echinoids 

occurring towards the northern half of the east Florida reef tract. To the north of Palm Beach County, 

nearshore reefs can have fairly high densities of the rock-boring urchins Echinometra lucunter

(Figure 4.19) and Arbacia punctulata. McCarthy 

(2001 and unpublished data) observed densities 

varying between 2 and 16 urchins/m2

 at several 

locations off Vero Beach. Approximately 

80% were E. lucunter, 19% A. punctulata, and 

1% the pencil urchin Eucidaris tribuloides. 

Echinometra lucunter also is found in tidal pools 

in the intertidal and in the very shallow subtidal 

zone along the entire coast. In this region, there 

may be an increase in abundance of A. 

punctulata with depth. This may be explained by 

the colder temperature tolerances of this species, 

as it is found much further north on the eastern 

coast than any of the other species (Hendler 

et al., 1995). The echinoid Tripneustes 

ventricosus also has been recorded as far north 

as Indian River County but generally occurs in 

low densities (C. Miller, Coastal Eco-Group, Inc., 

and D. McCarthy, Jacksonville University, 

Jacksonville, FL, pers. observ.). Progressing 

southward, in Martin County, several other species may be encountered on NHBs. These include 

Diadema antillarum, Lytechinus variegatus, and T. ventricosus. Of these three species, D. antillarum is 

more normally encountered on coral reefs and hardbottoms, whereas the other two are more common on 

seagrass habitats. D. antillarum have been observed throughout the east Florida coast but appear to be 

most abundant at the St. Lucie Reef (SFCREMP, 2007). Generally, south of Martin County, echinoid 

biomass decreases significantly, with only occasional observations of E. tribuloides and even rarer 

encounters with D. antillarum (D. McCarthy, 

Jacksonville University, Jacksonville, FL, pers. 

observ.). 

Very little is known about the holothurians along the 

Florida coast. The most conspicuous sea 

cucumbers are holothuroids. Holothuria grisea is 

most abundant in the intertidal and shallow subtidal 

zones along the east Florida coast (Figure 4.20) 

(Hendler et al., 1995). It can be found within cracks 

and crevices within worm rock or associated 

hardbottoms. It appears to occur in highest 

densities from Martin County and northward, 

although this has not been experimentally 

confirmed. The diversity of holothurians probably 

increases with depth. J. Beal (FWC, pers. comm.) 

identified five holothurians at the St. Lucie Reef in 

Martin County. 

Very little is known about the brittle stars and 

crinoids that occur along the east coast. There are 

probably a number of brittle star species that may be 

in high abundance along the coast, as they are somewhat easily encountered upon dissection of worm 

rock (D. McCarthy, Jacksonville University, Jacksonville, FL, pers. observ.). They also can be observed 

Figure 4.19. The rock-boring urchin Echinometra 

lucunter on shallow hardbottom 

habitat in Vero Beach, Florida. 

Photo by D. McCarthy.

Figure 4.20. The sea cucumber Holothuria grisea

on shallow hardbottom habitat in 

Ambersands Beach, Florida. 

Photo by D. McCarthy.
Section 2 4-35 

Organismal Assemblages of East Florida Shallow Hardbottom 

in the deeper subtidal hardbottoms from Palm Beach County southward. In contrast, crinoids probably 

are not that abundant but have been encountered in some areas. For instance, the crinoids Davidaster 

rubiginosa and Nemaster grandis have been identified in a depth of ~6 m at the St. Lucie Reef (J. Beal, 

FWC, pers. comm.). 

4.3.3.3 Reproduction and Life History 

All of the echinoderm species found along east Florida reproduce sexually. Most are slow egg producers 

(Eckelbarger, 1994) that seasonally spawn. While some do brood their young and have crawl-away 

larvae, many broadcast spawn, where fertilization results in either planktotrophic or lecithotrophic larvae. 

Larval time spent in the plankton varies from a few hours to months. After settlement and 

metamorphosis, many probably live for several years (Ruppert et al., 2004). 

While there is some information available on seasonal spawning patterns of echinoderms along the east 

Florida coast, there is little known on their dispersal and recruitment patterns. E. lucunter probably 

spawns during late summer/early fall, whereas A. punctulata spawns during late spring/early summer 

(Serafy, 1979). L. variegatus has two spawning periods---one during late spring and a second smaller 

peak in fall (McCarthy and Young, 2002). D. antillarum spawning periods appear somewhat variable 

throughout the Caribbean populations studied but often have a well-defined lunar rhythm (Hendler et al., 

1995). At Walton Rocks, Fort Pierce, the sea cucumber H. grisea occurs in low abundance during winter, 

when the animals may migrate offshore (Hendler et al., 1995). While most echinoderms found in our area 

produce planktonic larvae that may spend several months in the plankton before settling and recruiting to 

hardbottoms, there may be high dispersal and connectivity among echinoderm populations in the 

Florida-Caribbean region. There is no substantial research that has documented recruitment patterns for 

any echinoderm species along the east Florida coast. However, studies of echinoderms generally reveal 

recruitment to be low, if not sporadic, during the course of a single year. 

4.4 SUMMARY 

4.4.1 Diversity 

There is a high diversity and abundance of sessile and motile invertebrate species associated with NHBs 

along the east Florida coast (Appendix B, Table B.1). A total of 534 species of invertebrates has been 

identified along the east Florida coast, and there are likely to be more. The diversity of sessile species is 

greatest for cnidarians (~21%), bryozoans (~29%), and sponges (~19%). At generally lower diversities, 

tunicates, polychaetes, and bivalves are commonly encountered in these habitats as well. In contrast, 

some sessile taxonomic groups may not be represented with as high a diversity but can occur in very 

high biomass. A premier example is the sabellariid polychaete P. lapidosa, which can be very abundant 

along the more mid- to northern sections of the Florida coast. 

The most diverse and generally abundant motile invertebrates are arthropods (~47%) and polychaetes 

(~28%). There are over 100 species of crustaceans that are extremely abundant, especially on 

worm reef-dominated hardbottoms. These include crabs, stomatopods, shrimp, lobsters, isopods, and 

amphipods. There are over 87 reported polychaete species that are likely to be very abundant, although 

more research is needed to confirm this. There are also fairly high numbers of gastropods, flat worms, 

ribbon worms, and echinoderms on these habitats. While less diverse, to the north along the east Florida 

coast there is generally high biomass of echinoderms such as the sea urchins E. lucunter and 

A. punctulata. While these are some generally-observed trends, some groups such as brittle stars are 

probably fairly diverse and abundant but have not been extensively studied. 

4.4.2 Trophic Patterns and Functional Groups 

The two premier ecological functional roles of invertebrates along the east Florida coast are as 

1) shelter-enhancing organisms that increase local diversity of fishes and invertebrates, and 2) either 

predators or prey in local food webs. Generally, the highest community biomasses along the east Florida 

coast occur in hardbottom areas with higher abundances of sessile invertebrate species (some may be 
Section 2 4-36 

Organismal Assemblages of East Florida Shallow Hardbottom 

considered foundational or keystone contributors to the community) that enhance local shelter. Along the 

Florida coast, important shelter-enhancing taxonomic groups are hard and soft corals, sponges, 

tunicates, molluscs, barnacles, and polychaetes (i.e., P. lapidosa), however, their importance in this 

function varies dramatically with depth and latitude. In terms of contribution to local food webs, important 

taxonomic groups are sponges, crabs, shrimp, polychaetes, echinoderms, crabs, and shrimp. Again, 

their importance in this function likely varies dramatically with depth and latitude. 

The range of trophic roles is very different between sessile and motile east Florida invertebrates. Most 

sessile invertebrates are passive consumers, either being suspension feeders (sponges, sabellariid 

polychaetes, barnacles, tunicates) or opportunistic carnivores (hard and soft corals, bryozoans, 

hydrozoans). All of these invertebrates may be fed on by motile invertebrates or fishes, but there is little 

information available on their role in this function. In contrast, there is more trophic diversity among motile 

invertebrates. Most decapods and gastropods are active carnivores, although some are omnivores, 

herbivores, and suspension feeders. Echinoids such as E. lucunter, A. punctulata, and D. antillarum are 

important herbivores in the areas where they occur. These motile groups are also likely to be fed on by 

motile invertebrates or fishes, but there is little information available on their role in this function. 

4.4.3 Latitudinal and Cross-shelf Distribution 

On a large scale, the diversity and abundance of invertebrates along the east Florida coast varies 

dramatically based on latitude, depth, and, often, season. Variability in these community and population 

measures can be very high because of changes in features of the substrate itself, local hydrodynamics, 

and other biotic and abiotic factors. However, while there is not a good quantitative comparison of 

community assemblages along the coast, there are some general trends that appear to occur. 

Sessile invertebrate communities change noticeably with latitude and depth along the Florida coast. In 

intertidal hardbottoms to the north, dominant invertebrates are bryozoans, sabellariid polychaetes, sea 

anemones, and tunicates, although they generally do not occur in high biomass. In the more 

mid-sections of the east Florida coast, there is generally an increase in abundance of P. lapidosa, both 

intertidally and subtidally to approximately 4 m depth. Throughout tidal pools in these intertidal areas, 

Siderastrea spp., two species of zoanthids, and several species of solitary anemones can be 

encountered. Within the lower intertidal and shallow subtidal zones, the most conspicuous sponge is the 

rock-boring sponge Pione (formerly Cliona) lampa. In the more northern counties (Brevard and Indian 

River Counties) that have subtidal hardbottom, sponges, two species of gorgonians (Leptogorgia virgulata

and Leptogorgia hebes), and one species of telestacean (Carijoa (Telesto) riisei) can be encountered. In 

this area, the most significant hard corals are Siderastrea spp. and the Oculina species. The latter two 

species become fairly abundant in excess of 6 m (Reed et al., 1982; D. McCarthy, Jacksonville University, 

Jacksonville, FL, pers. observ.). However, there are occasional occurrences of Cladocora arbuscula and 

grouped polyps of Phyllangia americana and Astrangia spp. (Brooke and Young, 2005; CSA International, 

Inc., 2008; Coastal Eco-Group Inc., 2008, 2009). Progressing southward along the coast, the diversity 

and abundance of anthozoans, particularly scleractinians, increases on subtidal hardbottoms (SFCREMP, 

2007; J. Beal, FWC, pers. comm.). Within Martin County, the St. Lucie Inlet Reef appears to be the 

northernmost extent of several of the reef-building corals that are important biodiversity contributors within 

the Caribbean region. Within the 5 to 8 m depth range, mean percent cover of sponges is highly variable 

in samples, ranging between 0.14% and 10.29% (SFCREMP, 2007). In Broward County, a large 

percentage of the coral community in the excess of 3 to 7 m depth range is composed of the scleractinian 

Acropora cervicornis (Moyer et al., 2003; Vargas-Angel et al., 2003; SFCREMP, 2007). 

4.4.4 Reproduction and Life History 

A wide range of reproductive strategies is seen among the invertebrates encountered along the east 

Florida coast NHB. Most sessile organisms can reproduce asexually and sexually, while the majority of 

motile invertebrates are sexual reproducers. Both sessile and motile invertebrates tend to be short-lived 

(particularly in intertidal and very shallow subtidal waters), with a capability to reproduce continuously or 

semi-continuously. 
Section 2 4-37 

Organismal Assemblages of East Florida Shallow Hardbottom 

Almost all sessile invertebrates can reproduce asexually and sexually, although the relative importance of 

these modes in maintaining the population is often unclear. Sponges, cnidarians, bryozoans, and colonial 

tunicates are all important shelter-enhancing Florida invertebrates that have these reproductive 

capabilities. In some cases, such as with the scleractinian A. cervicornis, it is known that they primarily 

proliferate via asexual reproduction. Asexual reproduction may be very important for a number of 

shelter-enhancing invertebrate species in the shallower hardbottoms, where there is generally high 

disturbance. Noticeable sessile invertebrates that reproduce primarily via sexual reproduction are mostly 

polychaetes, bivalves, gastropods, and echinoderms. In particular, the reef-building polychaete 

P. lapidosa reproduces entirely via sexual reproduction, as do the vermetid gastropods in the genus 

Petaloconchus. 

Sexually reproducing sessile and motile invertebrates generally have seasonal spawning patterns, 

although many have the capability to spawn throughout most of the year. Seasonal spawning peaks 

often occur during the spring or summer time period for most of the invertebrates along the coast. 

However, the ability to continuously spawn is perceived to be an adaptation to take advantage of space 

made available by natural disturbances (McCarthy, 2001). For most sessile invertebrates, spawning is 

external and results in planktonic larvae, although there are instances of brooding and crawl-away young. 

4.4.5 Dispersal and Connectivity 

There is generally very little known regarding dispersal and connectivity of invertebrates along the east 

Florida coast. Considering known general reproductive capabilities of the various taxonomic groups, 

there is likely to be fairly low dispersal for many species, particularly the sessile ones. Tunicates, 

hydrozoans, some gastropods, amphipods, and isopods are likely to have the most limited dispersal 

capability. Sponges and hard and soft corals may be at an intermediate level of dispersal, while 

crustaceans, polychaetes, echinoderms, and bivalves generally should have higher dispersal and 

connectivity. However, several studies suggest that even some of these groups (i.e., Phragmatopoma, 

Acropora) may have fairly limited dispersal. 

4.4.6 Recruitment and Cross-shelf Habitat Use 

There is very limited information on seasonal recruitment patterns for invertebrates along the east Florida 

coast. There is essentially no information on cross-shelf habitat use. McCarthy (2001) studied the 

recruitment of sessile organisms in the intertidal and subtidal (3 to 4 m) zones at Boynton Beach, Florida 

from June 1997 to January 2000. He reported 23 species of sessile organisms recruiting to his 

settlement plates, with P. lapidosa being most commonly encountered (Table 4.1). The recruitment rate 

of some 7 to 11 species m2

/day occurred in both zones and was highest in summer and early fall. 

Although recruitment generally occurred at both depth zones, it was higher in the subtidal zone compared 

to the intertidal zone. McCarthy (2001) suggested that while high numbers of species may recruit into 

these habitats, most probably do not survive to adulthood. This may be due to a combination of factors. 

First, there was likely high mortality of recruits that could not tolerate physical factors that were very 

frequent, such as sand scouring and smothering. Second, high rates of P. lapidosa recruitment were 

observed to limit growth of sessile organisms, often growing over them. Finally, he also suggested that 

predation may not be as important in explaining recruit mortality in these two zones as the two 

previously-mentioned reasons. Overall, he suggested that worm reef-dominated hardbottoms appear to 

go through predictable patterns of annual change. These changes include high recruitment in early 

autumn through winter, rapid reef growth (approximately 0.5 cm/day) resulting in maximum structure in 

spring and summer, and decay by early autumn (Figure 4.9). When these data are integrated with those 

of Lindeman (1997a), they reveal important links between the seasonal cycle of sabellariid reef expansion 

and degradation and the occupation of those reefs by juvenile and adult organisms. 
Section 2 4-38 

Organismal Assemblages of East Florida Shallow Hardbottom 

4.4.7 Economic and Recreational Value 

Invertebrates significantly contribute to the economic and recreational value of NHBs along the east 

Florida coast. Sessile invertebrates such as P. lapidosa and a number of coral and sponge species 

provide high aesthetic value to NHBs as well as important shelter to many invertebrate and fish species of 

commercial and recreational value. Many of these species simply would not exist were it not for these 

foundational invertebrate species. Invertebrates also contribute to nearshore value via their significant 

role in the food web that again likely supports many commercially- and recreationally-important fish 

species. Finally, some invertebrates (e.g., spiny lobster) add value to the habitat as they are of direct 

commercial and recreational interest to the public. 

4.4.8 The Importance of Hardbottom to Invertebrates 

The presence of hardbottom is absolutely necessary for most of the invertebrates encountered along the 

east Florida coast. Sessile invertebrates such as corals, sponges, polychaetes, bryozoans, and tunicates 

are simply not found in neighboring sand bottom areas. Of these species, the lack of community 

contributors such as sabellariids and corals would likely result in the absence of most motile invertebrate 

and many fish species. 
Section 2 5-1 

Organismal Assemblages of East Florida Shallow Hardbottom 

Chapter 5

Fish Assemblages

5.1 INTRODUCTION 

NHB fish assemblages of mainland Florida can vary in diversity and total abundances. Very little journal 

literature exists on these assemblages. However, enough information is available from the gray literature 

and the scant research literature to allow fundamental evaluation of these often discrete, narrow habitat 

patches along east Florida that aggregate shallow reef organisms. 

Gilmore (1977) and Gilmore et al. (1981) discussed surf zone reefs and offshore reefs in terms of fishes 

and habitat features. In surf zone reefs in depths of 0 to 2 m, 105 fish species were documented. A total 

of 192 fish species from NHB areas within a depth range of 0 to 4 m was compiled from new data or the 

literature (Lindeman, 1997a). The number has increased (Lindeman and Snyder, 1999; Baron et al., 

2004) but has not been comprehensively revisited. Despite approximately three decades of monitoring of 

the juvenile-dominated and sometimes speciose NHB habitat (Figure 5.1), total species richness 

estimation is constrained by the paucity of published fish research in this habitat area. In addition, the 

existing fish species numbers may be low due to an absence of research on the diminutive, cryptic fish 

fauna, including gobiids, blenniids, and labrisomids. Ackerman and Bellwood (2000) and Robertson and 

Smith-Vaniz (2008) identified several problems regarding the underestimation of species diversity when 

cryptic ichthyofauna are chronically undersampled. 

Figure 5.1. Multispecies school of juvenile fishes including four species and three families 

(grunts, wrasses, and damselfishes) in association with nearshore hardbottom structure. 

Photo by D. Snyder.
Section 2 5-2 

Organismal Assemblages of East Florida Shallow Hardbottom 

Of the species recorded from NHB, the great majority of abundance and frequency records involve 

approximately 20 to 30 species; however, assemblage composition varies latitudinally and seasonally. 

The species are largely within approximately 10 families and include labrids, pomacentrids, haemulids, 

and carangids as prominent components (Lindeman and Snyder, 1999; Baron et al., 2004). To deal with 

this diversity, we identified four focal group categories based on groupings that share substantial habitat 

use and trophic attributes (Table 5.1): 

Demersal Carnivores: A diverse array of demersal piscivores, invertivores, and planktivores including 

grunts (Haemulidae), snappers (Lutjanidae), wrasses (Labridae), squirrelfishes (Holocentridae), drums 

(Sciaenidae), cardinalfishes (Apogonidae), sweepers (Pempheridae), and over 15 other families, 

including some blenniids and labrisomids. Grunts typically are the most abundant and diverse fish fauna 

on NHB structure of mainland east Florida, with 11 species recorded and over 80% of total individual 

abundance in available studies. Since new larval recruits feed on plankton, grunt early life stages can 

also be abundant planktivores on subsets of the shallow reef. This group also includes notable nearshore 

fishery species as adults or during developmental periods before maturation. 

Cryptic Hardbottom Residents: The gobies, blennies, sand stargazers, clingfishes, and related forms 

(Blenniidae, Labrisomidae, Clinidae, Chaenopsidae, Dactyloscopidae, Gobiesocidae) include at least 

25 species recorded from shallow hardbottom. Almost all of the cryptic, attached fishes have specialized 

morphological adaptations for NHB residency and a structure-attached life history after settlement. 

Though they typically perch on hardbottom, these species are trophically diverse. The gobiids and some 

blennies are herbivorous, but notable and mobile carnivores, including several families of eels, are also 

within this group (Table 5.1). 

Coastal Pelagics: Includes transient, water-column dwelling piscivores such as jacks, mackerels (often 

in large schools), and sharks (Carangidae, Scombridae, and Carcharhinidae) and small, schooling bait 

species including the planktivorous herring, anchovies, scads, and mullets (Clupeidae, Engraulidae, 

Carangidae, and Mugilidae, respectively), approximately 25 species in total. 

Herbivores and Omnivores: Includes approximately 20 species of damselfishes (Pomacentridae), 

parrotfishes (Scaridae), doctorfishes (Acanthuridae), chubs (Kyphosidae), porgies (Sparidae), and some 

species of filefishes (Monacanthidae) and spiny puffers (Diodontidae). The site-associated blennies 

(Blenniidae) are also herbivorous. Several of these species occur in very high abundance and may be of 

trophic significance in the uptake and energetic conversion of macroalgae and other items. 

Table 5.1. Focal groups of fishes based on trophic and habitat use attributes. 

Groups Approximate Number of 

Families and Species Example Families and Genera 

Demersal 

carnivores 

30 families; 

10 species 

Haemulidae (grunts, Haemulon and Anisotremus); Labridae 

(wrasses, Halichoeres); Sciaenidae (drums, Pareques and

Bairdiella); Lutjanidae (snappers, Lutjanus) 

Cryptic 

hardbottom 

residents 

8 families; 

30 species 

Gobiidae (gobies, Coryphopterus); Labrisomidae (scaled blennies, 

Labrisomus); Apogonidae (cardinalfishes, Apogon); Muraenidae 

(moray eels, Muraena) 

Coastal pelagics 10 families; 

25 species 

Carangidae (jacks, Caranx); Clupeidae (herring, Harengula); 

Scombridae (mackerels, Scomberomorus maculatus) 

Herbivores 7 families; 

20 species 

Pomacentridae (damselfishes, Stegastes and Abudefduf); 

Kyphosidae (chubs, Kyphosus); Scaridae (parrotfishes, Sparisoma); 

Acanthuridae (surgeonfishes, Acanthurus); Sparidae (porgies; 

Diplodus) 

For each focal group, prominent biological features are summarized with emphasis on spawning 

activities, shelter use characteristics, and trophic functioning in association with NHB, IHB, and shallow 

OHB. We do not refer to these as guilds; these groups are too diffuse and are used solely to simplify the 

taxonomic and ecological complexity of the fishes examined. Guilds often attempt to separate species 
Section 2 5-3 

Organismal Assemblages of East Florida Shallow Hardbottom 

groups according to only one primary functional category (e.g., feeding guilds or habitat associations 

[Appeldoorn and Lindeman, 1985; Elliot et al., 2007]), and we are not that restrictive in this case. 

An additional fish assemblage occurring largely over adjacent sand plain habitats, the surf zone fish 

assemblage, can co-occur with NHB and IHB structure and organisms, e.g., via trans-habitat feeding 

migrations. There is a small literature on the species characteristics of shallow sand plain and surf zone 

habitats of east Florida, including Gilmore et al. (1981), Peters and Nelson (1987), and Gorecki (2007). 

There are significant differences in the taxonomic and ecological species compositions of adjacent surf 

zone and NHB fish assemblages in east Florida. For example, different sciaenid faunas (croakers and 

drums) occur between NHB and sand plain areas (Gilmore et al., 1981). For the family Sciaenidae, the 

most ecologically and economically significant species associate with sand habitats more often than reef 

habitats. There are also important species and process interactions between surf zone and NHB fishes 

(e.g., in some sites, Spanish mackerel and pompano are often caught by fishers over NHB or near the 

edges of NHB and sand). Ontogenetic foraging events, particularly from NHB habitats over adjacent 

sand plain habitats, may become important as cohorts of hundreds of surviving and maturing juveniles of 

grunt, wrasse, and other species migrate increasingly away from the NHB to seek more prey and habitats 

associated with mature stages (Randall, 1967; Parrish, 1989), often but not always in deeper water. 

Based on the demonstrated or potential value of NHB areas as both nurseries and spawning sites for 

economically valuable species, these habitats were determined to support highly important ecological 

functions; a primary NMFS criterion for EFH-HAPC designation under the federal Sustainable Fisheries 

Act (1996). The mandate of the Sustainable Fisheries Act to conserve significant habitats drove 

substantial convergences between fishery management and habitat management in some areas of the 

country (SAFMC, 1998; Fluharty, 2000; Lindeman et al., 2000). The Federal EFH-HAPC designation has 

administrative significance for all species of haemulids, lutjanids, and serranids managed under the 

SAFMC Snapper Grouper Fishery Management Plan; this includes over 40 species, many of which are 

significant components of Florida’s 

approximately $6 billion per annum 

recreational salt water fisheries. At least 

15 of these managed species occur on NHB, 

typically as juveniles or earlier life stages. 

5.2 DEMERSAL CARNIVORES 

5.2.1 Focal Families and Species 

This category includes demersal invertivores, 

piscivores, and planktivores that are not 

site-attached to hardbottom structure 

(Table 5.1). Many families are represented 

in NHB habitats, with some species also 

having significant roles on deeper and 

structurally different hardbottom habitats 

after emigrating following settlement and 

maturation (Figure 5.2). Families include 

the grunts (Haemulidae, 11 species recorded 

from NHB), wrasses (Labridae, 11 species), 

snappers (Lutjanidae, 6 species), 

squirrelfishes (Holocentridae, 3 species), 

drums (Sciaenidae, 6 species), groupers 

(Serranidae, 9 species), threadfins 

(Polydactylidae, 2 species), sweepers 

(Pempheridae, 1 species), lizardfishes 

(Synodontidae, 3 species), and over 15 other 

families. 

Figure 5.2. Early juvenile lane snapper (Lutjanus 

synagris) associating with algal-rock-sand 

interface, 3 to 4 m, John U. Lloyd State 

Park, Broward County, Florida. Note rapid 

metamorphosis compared to haemulids. 

Photo by L. Jordan.
Section 2 5-4 

Organismal Assemblages of East Florida Shallow Hardbottom 

Over 80% of the individual abundances in the NHB fish assemblage in both northern Palm Beach County 

and central Broward County were represented by species of grunts and margates (Lindeman and Snyder, 

1999; Baron et al., 2004). Diverse and widely distributed, the 18 species of the genera Haemulon and 

Anisotremus (Haemulidae, Percoidei) comprise one of the most ecologically and economically important 

groups of reef fishes in the Western Atlantic (15 and 3 species, respectively) due to their high abundance 

and trophic importance as predators and prey across diverse cross-shelf habitats. There are 12 haemulid 

genera in the Western Hemisphere, and these taxa occupy a spectrum of habitats from riverine to deep 

reef (Lindeman and Toxey, 2003). Many of the reef species typically use nearshore habitats during 

settlement and early juvenile maturation periods and utilize offshore reefs during older life stages. 

Some demersal carnivore species are significant in recreational or commercial fisheries. Their 

species-specific usage patterns of NHB may range from 1) habitat use primarily during early life stages, to 

2) adults using NHB or adjacent areas for feeding. These species include snappers and groupers in the 

former category and pompano (Trachinotus carolinus) and sheepshead (Archosargus probatocephalus) 

in the latter group. Commonly under-recognized as NHB users, some of the most sought-after inshore 

gamefish species are caught on NHB habitats throughout several NHB areas of the project region, 

including tarpon (Megalops atlanticus), red 

drum (Sciaenops ocellatus), snook 

(Centropomus undecimalis), and sea trout 

(Cynoscion spp.), the majority capable of 

feeding on both fishes and invertebrates. 

Larger demersal carnivores of shallow 

hardbottom include sharks (Figure 5.3). 

One demersal carnivore, striped croaker 

(Bairdiella sanctaeluciae), is listed as a 

species of special concern by state and 

federal agencies because its distribution in 

U.S. waters is limited to eastern Florida. 

This species occurs over NHB and OHB 

from Jupiter to Melbourne, Florida and can 

be locally abundant on NHB (Gilmore and 

Snelson, 1992) and shallow water artificial 

reefs (Continental Shelf Associates, Inc., 

2006; Coastal Eco-Group Inc., 2009). 

5.2.2 Shelter Use 

5.2.2.1 Spawning Sites and Larval Transport 

Spawning in many fish families that use shallow hardbottom is still poorly known. Some of the species in 

this category spawn in aggregations, but many spawn in pairs (Thresher, 1984). Snapper species 

(Lutjanidae) form spawning aggregations in southern Florida and Cuba that can potentially generate 

propagules that settle in eastern mainland Florida (Lindeman et al., 2000; Paris et al., 2005), but they do 

not spawn directly on shallow hardbottom. On the other end of the life history scale are families that 

greatly limit their dispersal such as jawfishes (Opistognathidae), which brood eggs in their mouths. 

Predicatively understanding local and meso-scale physical oceanography as well as behavioral 

processes will be critical to addressing such dispersal issues further. Whether the species has life history 

traits that predispose it to wide dispersal or local retention, larval transport is also poorly known in terms 

of empirical studies. Some useful information is available from studies focusing on receipt in east Florida 

of upstream snapper larvae from known snapper spawning aggregations in Cuba. Coupled biophysical 

modeling for several demersal snapper species found on east Florida NHB suggests advection rates of 

lutjanid larvae (lane snapper, Lutjanus synagris, and gray snapper, L. griseus) from Cuba to east Florida 

are relatively low (Paris et al., 2005). 

Figure 5.3. Resting nurse shark, Ginglymostoma cirratum, 

a generalized feeder on bottom invertebrates, 

3 m depth, Palm Beach, Florida. 

Photo by D. Snyder.
Section 2 5-5 

Organismal Assemblages of East Florida Shallow Hardbottom 

Despite their abundance, information on reproduction in grunts is limited (García-Cagide et al., 2001). 

Fishermen have reported potential spawning aggregations, but this information requires validation (Claro 

and Lindeman, 2003), and spawning has not been observed in any Western Atlantic haemulids. There is 

little evidence that any species of Haemulon would spawn in association with NHB. Black margate could 

be an exception in deeper NHB or IHB areas, especially near channels, since this species can grow to 

maturity in very shallow water and there is indirect evidence that it may spawn in nearshore channels. 

The potential spawning of other haemulid species in deeper IHB or shallow OHB is not precluded, 

especially given the unusual early life history details discussed below. 

Considerable evidence suggests that larvae of Haemulon and Anisotremus do not commonly leave 

insular or continental shelves, may associate with the bottom in transient manners within a week after 

hatching, and undergo a slow epibenthic metamorphosis that extends well past settlement (Lindeman et 

al., 2001). The evidence includes post-settlement planktivory (Helfman et al., 1982), a slow 

morphological metamorphosis away from larval characteristics (translucence, incomplete fin 

development) that continues for weeks post-settlement (Lindeman, 1986; Lindeman and Richards, 2005), 

and the absence of haemulid larvae from offshore ichthyoplankton surveys (Richards, 1984; Sponaugle et 

al., 2003). This suggests that larval dispersal may be constrained by behaviors and positioning that 

appear to not favor advection by flow fields. 

Retention on the shelf in shallow water coupled with such early settlement, possibly involving multiple 

bottom-association events prior to conventional “settlement,” is a relatively uncommon early life history 

strategy (Lindeman et al., 2001). The typical settlement pattern for many percoid and other important 

groups of families involves direct settlement with a rapid metamorphosis (one to several days). Grunts, a 

basal-intermediate percoid family, have a slower and different demersal metamorphosis process from 

many other species and families. 

5.2.2.2 Settlement and Early Associations with Shelter 

Settlement and early habitat use of the species in the demersal carnivore group are not well known; 

however, general patterns that characterize certain taxa provide some insight. Most groupers are very 

secretive at settlement and are rarely observed during visual surveys. Another characteristic of groupers 

and several other species including snappers is settlement on inshore seagrass meadows. Individuals of 

these species will, as they grow, migrate to NHB habitats. Lane snapper and mahogany snapper 

(Lutjanus mahogoni) are two lutjanid species that appear to settle directly on NHB (lane snapper also will 

settle in seagrass meadows). 

Grunts and margates (Haemulon and Anisotremus species) settle by 7 to 10 mm standard length (SL) at 

the least. These are small settlement sizes compared to most reef fish families, and grunts possess an 

unusual epibenthic larval stage as part of a highly extended metamorphosis that proceeds well past 

settlement (Figure 5.4). Understanding of the dense newly settled schools that grunts can form has been 

complicated by a sequence of two post-settlement pigment patterns (newly settled and early juvenile) that 

most species possess during an extended epibenthic metamorphosis (Lindeman, 1986; Lindeman et al., 

2001) (Figure 5.5). 

Few speciose genera of Western Atlantic reef fishes show such extended morphological and ecological 

metamorphoses at settlement. The newly settled pattern between 7 and 15 mm SL shows transitional 

attributes of both larval and early juvenile pigmentation and continuation of planktivory into 

post-settlement stages (Figure 5.4); delayed metamorphoses are not present in larval recruits of many 

other percoid fishes. Grunts are among the most common settlers in many differing shallow coastal 

habitats (McFarland et al., 1985; Shulman and Ogden, 1987; Lindeman and Snyder, 1999; Baron et al., 

2004) and are the most common settler on shallow hardbottom of east mainland Florida of all the families 

in the huge perciform suborder Percoidei. Evolutionarily, the only families that are similarly abundant are 

in the very different suborder Labroidei (e.g., damselfishes and wrasses) (Kaufman and Liem, 1982). 
Section 2 5-6 

Organismal Assemblages of East Florida Shallow Hardbottom 

Figure 5.4. Newly settled grunts in the transitional epibenthic planktivore stage (approximately 12 to 

20 mm standard length) after settlement and before adoption of demersal feeding (by the 

late early juvenile stage), Palm Beach, Palm Beach County, Florida. Photo by D. Snyder.

Figure 5.5. Early life stages of primary reef-associating genera of haemulids and lutjanids of the western 

Atlantic. Life stage designations and size ranges (mm standard length) based on 

morphological and ecological information from Starck, 1970; Lindeman, 1986; Richards, 

1994; and Lindeman and Richards, 2005. Representative figures are of newly settled and 

early juvenile stages of smallmouth grunt (Haemulon chrysargyreum) and mutton snapper 

(Lutjanus analis). 
Section 2 5-7 

Organismal Assemblages of East Florida Shallow Hardbottom 

Sweepers (Pempheris schomburgkii) and drums (Pareques spp., Bairdiella sanctaeluciae, and 

Odontoscion dentex), like the grunts and margates, settle in groups near vertical ledges where they are 

readily observed. The common wrasses such as slippery dick, puddingwife (Halichoeres radiatus), and 

clown wrasse may exhibit a very different pattern of settlement: presumably settling and burying 

themselves in sand bottom, remaining there for at least 5 days before emerging to take up residence on 

the reef (Victor, 1991). 

5.2.2.3 Juvenile and Adult Habitat Use 

On hardbottom from 0 to 7 m deep (mean depth of 3.9 m) in Broward County, 84% of all fishes and over 

90% of all juvenile fishes surveyed were species of Haemulon (Baron et al., 2004). This study covered 

areas with wide eastward hardbottom strata crossing among NHB, IHB, and shallow OHB depth zones, in 

contrast to many NHB areas in northern Palm Beach County. Four of the top six most abundant species 

in transects were grunts, along with H. bivittatus and Stegastes variabilis (Baron et al., 2004). The most 

abundant grunt was “Unidentified Haemulon sp.” with over 8,000 individuals, with a mean total length (TL) 

of 2.6 cm for transect counts. 

In surveys at Coral Cove County Park and Carlin Park in northern Palm Beach County, and Ocean Ridge 

in southern Palm Beach County, 7 of the top 12 most abundant species from the three sites were grunts 

and margates, in order: sailors choice (Haemulon parra), black margate (Anisotremus surinamensis), 

porkfish (A. virginicus), tomtate (H. aurolineatum), unidentified Haemulon spp., French grunt 

(H. flavolineatum), and smallmouth grunt (H. chrysargyreum). In all species, the most abundant life 

stages were early juvenile and juvenile stages, newly settled stages, and adults. Adult abundances were 

significantly different from pooled early life history stages in all species tested (Figure 5.6). 

Figure 5.6. Comparisons of abundances of life history stages at two sites in northern Palm Beach 

County (From: Lindeman and Snyder, 1999). Boxes note comparisons between adult 

stages and all sub-adult stages pooled. A: All species pooled. B: Only grunts (species of 

Haemulon and Anisotremus) pooled. NS = newly settled; EJ = early juvenile; J = juvenile; 

PE = pooled early stages. 
Section 2 5-8 

Organismal Assemblages of East Florida Shallow Hardbottom 

5.2.3 Trophic Patterns 

There are no fish feeding studies from shallow hardbottom habitats in the project region. The most 

detailed examination of food habits of Caribbean reef and coastal pelagic fishes is Randall (1967). His 

study areas were in the U.S. Virgin Islands and Puerto Rico. These are insular systems with both 

similarities and differences in the seascapes that these animals move across compared to east Florida. 

The grunts and margates of the genera Haemulon and Anisotremus show a wide array of invertivory. 

Some species are planktivorous as adults (H. striatum), others are very specialized for feeding on 

hard-shelled invertebrates (H. macrostomum), while the majority of species are relatively generalized 

feeders on both soft- and hard-invertebrate prey, often from either softbottom or hardbottom 

environments (Randall, 1967; Sierra, 2001). 

Most species are primarily demersal invertivores, as well as some important demersal piscivores and 

planktivores. Over 50 species of NHB fishes consume invertebrates as a primary dietary component. 

Eleven species of wrasse illustrate a pattern similar to grunts and other families: invertebrates from many 

diverse taxa are consumed by differing congeners with low to high levels of specialization within the 

genus. For example, many wrasse species (genus Halichoeres) feed on “shelled invertebrates” in the 

northeast Caribbean, though specific dietary compositions can vary (Randall, 1967; Randall’s data 

tabulated by taxa in Sierra, 2001). The dominant labrid in both the Jupiter and Fort Lauderdale studies 

was the slippery dick (H. bivittatus). This species feeds on crabs, urchins, annelids, and gastropods 

(Randall, 1967) and is an opportunistic feeder. Other labrids, such as the clown wrasse, H. maculipinna, 

feed primarily on soft annelids (Randall, 1967). As mentioned elsewhere, these comparative patterns are 

largely predictive without empirical feeding information from the project region. 

Additional invertivores on NHB habitats include snake eels (Ophichthidae), one species of moray eel 

(chain moray, Echidna catenata), and squirrelfishes (Holocentridae), which feed on crabs, shrimp, and 

molluscs. Smaller serranids (Serranus spp.) and reef drums including Pareques lanceolatus and 

P. punctatus prey upon shrimp, echinoids, and annelids. Several species of goatfishes (Mullidae) and 

some labrisomid species prey upon swimming crabs and ophiuroids. Juvenile stages of most snapper 

species feed on crabs and shrimp and with growth switch to fishes. The boxfishes (Ostraciidae) broadly 

feed on sponges, sipunculids, tunicates, and crustaceans. Spiny puffers (Diodontidae) feed on 

gastropods, echinoids, and crabs (Randall, 1967). 

Piscivorous families on NHB include the lizardfishes (Synodontidae) and older groupers and snappers. 

Most species of moray eels are also piscivores, but these are considered under cryptic hardbottom 

residents. These species consume a high diversity and abundance of smaller prey fishes above or 

adjacent to reef structure. Common prey items include juvenile stages of grunts, wrasses, and herring 

(Randall, 1967). 

Demersal planktivory by older juveniles and adults is present among cardinalfishes (Apogonidae), 

sweepers (Pempheridae), and species such as yellowtail snapper. The apogonids reside in crevices 

during the day and feed on the larvae of shrimp, crabs, and annelids in the northeast Caribbean (Randall, 

1967). Based also on the work of Randall (1967) and Gladfelter (1979), the copper sweeper (Pempheris 

schomburgkii) also resides near crevices and overhangs where it feeds on annelid larvae and other 

plankton. Large schools of newly settled individuals are commonly observed. They appear to possess 

an extended epibenthic larval period similar to grunts. Less common yellowtail snapper feed on a variety 

of plankton in the water column above the surface of hardbottom structure.

5.2.4 Developmental Patterns and Cross-shelf Habitat Use 

The great diversity of families and species of demersal invertivore groups and the absence of habitat use 

literature from east Florida NHB constrains broad summary, as does geographic variation. However, the 

available information suggests over 30 species of haemulids, labrids, lutjanids, sciaenids, serranids, and 

others typically redistribute across the shelf during development, often into deeper waters. 
Section 2 5-9 

Organismal Assemblages of East Florida Shallow Hardbottom 

The degree to which maturing juveniles on discrete NHB patches can successfully emigrate to deeper 

water reef structure is unexamined. Based on the data and evidence from other studies, some species 

assumedly should attempt to emigrate away at a certain age. We assume this process is gradual and 

involves exploratory behavior that may be associated with nocturnal sand plain feeding migrations in 

some species (e.g., snappers, grunts). The advent of small telemetric tags used in conjunction with a 

coordinated set of cross-shelf receiving stations could help resolve this issue for some species.

5.2.5 Latitudinal Distribution 

Information from multiple sites is limited and comparisons are complicated by variable relative 

abundances of NHB, IHB, and connected OHB habitats among the counties (Table 2.1), the very different 

geomorphologies and oceanographic conditions latitudinally, and by the differing depths examined in the 

existing studies. 

Some differences among ichthyofaunal assemblages are seemingly present between the southern and 

northern areas of mainland southeast and east-central Florida in terms of the most abundant species. 

Baron et al. (2004) found one black margate, Anisotremus surinamensis, out of >72,000 individuals from 

164 families recorded in Fort Lauderdale in the summer of 2001. To the north, Lindeman and Snyder 

(1999) found 636 A. surinamensis out of 10,491 individuals censused, 6% of the overall assemblage. 

A. surinamensis in the Jupiter area ranked seventh in individual abundance of over 80 species recorded. 

In the same studies, Baron et al. (2004) found 43 sailors choice, Haemulon parra, out of 72,000 fish 

recorded; most were older juveniles with mean lengths between 15 and 17.5 cm. A total of 58,000 grunts 

ranging from 1 to 7 cm TL (mean of approximately 2.6 cm), >80% of the total number of species 

recorded, was identified as Haemulon sp. in the Broward research. H. parra in the Jupiter area ranked 

first in abundance of all species. Lindeman and Snyder (1999) found 1,326 H. parra individuals, primarily 

early juvenile and newly settled stages, 13% of all fishes recorded in 2 years. There is a similar 

difference between south and north abundances in the hairy blenny, Labrisomus nuchipinnis; a dominant 

cryptic hardbottom resident species of the Jupiter area and sites to the north (Gilmore, 1977; Gilmore et 

al., 1981; Lindeman and Snyder, 1999) that was uncommon in Broward surveys (Baron et al., 2004). 

Given the high number of species and families and the substantial physiographic transitions between 

Miami and Cape Canaveral, there have to be some species whose distribution and abundance patterns 

consistently differ between the northern and southern areas of the project region. Absent further 

empirical information from more equivalent depth settings, the early comparisons of latitudinal variation 

above will remain somewhat tenuous until more studies become available. 

5.3 CRYPTIC HARDBOTTOM RESIDENTS 

5.3.1 Focal Families and Species 

Shelter association and trophic diversity are high among cryptic hardbottom resident species. For 

example, five primary cryptic resident families (gobiids, labrisomids, clinids, blenniids, and chaenopsids) 

and their over 25 phylogenetically similar species show feeding patterns that vary from herbivory (all 

gobiids and some common blennies) to invertivory (cardinalfishes, squirrelfishes) and piscivory (eels).

5.3.2 Shelter Use 

5.3.2.1 Spawning and Larval Transport 

Little is known of the spawning and larval biology of cryptic hardbottom resident species in east Florida. 

The available evidence suggests that most cryptic NHB-associated species can build nests. Compared 

to some other nest layers at these depths (e.g., damselfishes), the nests of most cryptic residents of 

NHB-IHB habitats are typically more cryptic, often under overhangs or deep in small holes and crevices. 

The molly miller (Scartella cristata) will deposit eggs in empty barnacle shells or abandoned worm holes 

(Breder and Rosen, 1966). Direct assessment of spawning biology is limited. Indirect evidence includes 
Section 2 5-10 

Organismal Assemblages of East Florida Shallow Hardbottom 

color patterns associated only with courtship and 

spawning (Figure 5.7). Gobies, blennies, and 

associated cryptic resident taxa tend nests of eggs. 

There is considerable information from some congeneric 

gobiids and blenniids from the Pacific coast of North 

America (or confamilials). In these species, often 

residing in tidepools and rocky subtidal and intertidal 

habitats, the available evidence suggests larval 

dispersal patterns are frequently limited (Gunderson et 

al., 2008). Larval retention is a primary early life history 

characteristic of many rocky intertidal species (Marliave, 

1986; Strathmann et al., 2002; Broitman et al., 2008; 

Gunderson et al., 2008). 

5.3.2.2 Settlement and Early Habitat 

Associations 

Early development and settlement is also poorly known 

in the cryptic residents of shallow hardbottom of east 

Florida. Breitburg (1991) documented features of 

demersal schooling before settlement in the naked goby. 

Developmental studies of many species detail how the pectoral fins are modified and fused into ventral, 

cup-like structures that allow direct attachment to NHB structure in high wave energy environments. 

These structures are formed by the time the age at which settlement can occur is reached.

5.3.2.3 Juvenile and Adult Habitat Use 

Cryptic resident species do not appear to migrate 

further offshore with development, displaying no 

cross-shelf ontogenetic migrations. They are 

life-long residents of the NHB, IHB, or shallow 

OHB structure on which they settle, and display 

feeding modes from herbivory to probable 

piscivory (Figure 5.8). Nine species of gobies and 

blennies were recorded as “restricted to the 

nearshore” in Baron et al. (2004): four labrisomid 

species, four gobiids, and one blenniid species; 

this number represents 45% of all the species they 

estimated to be nearshore-restricted. 

Hairy blenny was the largest and most abundant 

cryptic hardbottom resident in the Jupiter surveys, 

with 806 individuals recorded out of 

10,491 individuals censused, 8% of the overall 

individual total (Lindeman and Snyder, 1999). 

Hairy blenny ranked fifth in individual abundance of over 80 species recorded. Hairy blenny was also the 

most abundant cryptic hardbottom species on Ambersand mitigation reef surveys, Indian River County, 

representing 30% of the individuals censused on artificial reefs and natural nearshore hardbottom over 

3 years of annual monitoring (Coastal Eco-Group Inc., 2009). Baron et al. (2004) recorded no hairy 

blennies on transect and point-counts in Fort Lauderdale, though the species was recorded in qualitative 

roving diver surveys. They found the bridled goby, Coryphopterus glaucofraenum, and the rosy blenny, 

Malacoctenus macropus, at 2 to 5 cm size ranges to be the most abundant cryptic resident species. 

Other cryptic taxa observed on NHB include moray eels (Muranenidae), snake eels (Ophichthidae), 

soapfishes (Serranidae), and cardinalfishes (Apogonidae). Juveniles and adults from these groups are 

Figure 5.7. Spawning coloration in male 

hairy blenny (Labrisomus 

nuchipinnis), with female in 

background, 2 m depth, Pecks 

Lake Reef, southern Martin 

County, Florida. Photo by 

T. Gibson.

Figure 5.8. A cryptic hardbottom resident, 

spotted moray eel (Gymnothorax 

moringa), depth 2 m, MacArthur 

Beach State Park, Palm Beach 

County, Florida. Photo by D. Snyder.
Section 2 5-11 

Organismal Assemblages of East Florida Shallow Hardbottom 

generally solitary, dwelling under ledges and in crevices provided by NHB, IHB, or OHB. All of these taxa 

are predatory and generally leave their hiding places to forage at night. Baron et al. (2004) recorded 

spotted moray, purplemouth moray, and goldentail moray offshore Fort Lauderdale; spotted snake eel 

also occurred within their study area. In Jupiter, Lindeman and Snyder (1999) observed all of these eels 

except the purplemouth moray, but also reported the chain moray (Echidna catenata). 

Cardinalfishes (apogonids) were more diverse and abundant in visual samples recorded in Fort 

Lauderdale (Baron et al., 2004) in data from 0 to 7 m. Two genera and six species were reported, with 

twospot cardinalfish (Apogon pseudomaculatus) and flamefish (Apogon maculatus) represented by the 

most individuals. Cardinalfishes were less commonly seen in Jupiter censuses; flamefish was the only 

species recorded. Greater soapfish (Rypticus saponaceous) is the most commonly observed soapfish in 

the region and was infrequently seen by Baron et al. (2004) and Lindeman and Snyder (1999). 

5.3.3 Trophic Patterns 

Highly diverse feeding patterns exist among cryptic hardbottom fish species. Herbivory, invertivory, 

planktivory, and piscivory are all known for the small fishes that attach to NHB structure, often out of 

human sight. Randall (1967) found that “all gobiids” and two common blenny species (Scartella cristata

and Parablennius marmoreus) fed primarily on algae in the U.S. Virgin Islands and Puerto Rico. 

Labrisomids such as the very common hairy blenny, a generalized invertivore in Randall (1967), probably 

can also feed upon late stage larvae of some reef-associated fishes such as grunts (R.G. Gilmore, Jr., 

Estuarine, Coastal, and Ocean Science, Inc., Vero Beach, FL, pers. comm.). First-ever food habit studies 

on these and other NHB species would be of great value, as knowledge of the trophic structure of this 

east Florida fish subassemblage is depauperate. 

5.3.4 Latitudinal Distribution 

As was the case for grunts, there is a contrast between the southern and northern sites surveyed in 

Baron et al. (2004) and Lindeman and Snyder (1999). No hairy blennies were recorded on transect and 

point-counts in Fort Lauderdale, though the species was recorded in qualitative roving diver surveys 

(Baron et al., 2004). In contrast, hairy blenny was the largest and most abundant cryptic hardbottom 

resident in the Jupiter surveys to the north, with 806 individuals recorded out of 10,491 individuals 

censused, 8% of the overall individual total (Lindeman and Snyder, 1999). Hairy blenny ranked fifth in 

individual abundance of over 80 species recorded. 

By definition, the visual methods used by Baron et al. (2004) and Lindeman and Snyder (1999) will not 

fully account for all cryptic species present in the area. Accordingly, other cryptic taxa were seldom 

observed and therefore contribute little information to support analyses of latitudinal patterns. 

Nevertheless, some basic patterns may be described for the soapfishes and cardinalfishes. For 

soapfishes it appears that the whitespotted soapfish is more common north of Broward County and the 

greater soapfish is more prevalent from Palm Beach County southward. As mentioned above, the 

cardinalfishes were much more diverse in the southern area samples, and observations on OHB indicate 

that cardinalfish species drop out with increasing latitude to about Jupiter; from this area north, twospot 

cardinalfish and flamefish are the most commonly-observed species on OHB and NHB. 

5.4 COASTAL PELAGICS 

5.4.1 Focal Families and Species 

Though typically transient, coastal pelagic fish species can interact with NHB in manners that have local 

trophic impacts. Their roles as both predators and prey can be seasonally significant. We therefore 

consider coastal pelagic species as a transient subcomponent of the larger NHB-associated 

ichthyofauna. The species in the coastal pelagic functional group are water-column dwelling, often 

schooling fishes that are typically moving. These species usually do not reside on NHB for extended 

periods of time. 
Section 2 5-12 

Organismal Assemblages of East Florida Shallow Hardbottom 

The diverse taxa of the coastal pelagic group includes 

• small, schooling herring, anchovies, scads, mullets, and halfbeaks (Clupeidae, Engraulidae, 

Carangidae, Mugilidae, and Hemiramphidae, respectively), prey to many NHB resident and transient 

species; 

• large, schooling mackerels, bluefish, and pompanos (Scombridae, Pomatomidae, and Carangidae, 

respectively) in cooler months, predators on many nearshore fishes and prey to anglers; and 

• the largest apex marine predators of coastal Florida, several shark species such as nurse, requiem, 

and hammerhead (Ginglymostomatidae, Carcharhinidae, and Sphyrnidae, respectively), some of 

which are commonly solitary, whereas others can form schools. Many of these species, including 

blacktip (Carcharhinus limbatus), blacknose (C. acronotus), bull (C. leucas), spinner (C. brevipinna), 

tiger (Galeocerdo cuvier), and lemon (Negaprion brevirostris) sharks, are federally managed (NMFS, 

1999). 

5.4.2 Shelter Use 

5.4.2.1 Spawning and Larval Transport 

Some of these species have little or no association with particular bottom types while spawning. 

Structural components of the water column, such as temperature gradients, may have more of a habitat 

influence than bottom type (Kingsford et al., 2002). Several clupeids have been observed with 

developing or ripe ovaries in nearshore waters of Palm Beach County, particularly Spanish sardine 

(Sardinella aurita), false pilchard (Harengula clupeola), and scaled sardine (H. jaguana) (D. Snyder, CSA 

International, Inc., pers observ.). Bigeye scad (Selar crumenophthalmus) also has been collected with 

ripe ovaries in nearshore waters adjacent to NHB (Continental Shelf Associates, Inc., 1992). 

Although there is little direct evidence of coastal pelagic species spawning over or adjacent to NHB, when 

depth preferences and behavior are considered, it could occur in some species. For example, Atlantic 

thread herring (Opisthonema oglinum), Spanish sardine, and round scad (Decapterus punctatus) would 

probably spawn in deeper offshore waters, whereas the false pilchard, scaled sardine, and redear sardine 

(Harengula humeralis) are more likely to spawn in shallow habitats such as NHB (e.g., Martinez and 

Houde, 1975). Another group that spawns in shallow coastal waters and has been observed over NHB, 

IHB, and OHB off Palm Beach (the northern limit of its distributional range) are the halfbeaks 

(Hemiramphus brasiliensis) and (H. balao) (Berkeley and Houde, 1978; McBride et al., 2003). 

5.4.2.2 Settlement and Early Habitat Associations 

Many coastal pelagic species do not appear to possess a discrete, bottom-associated settlement event 

associated with NHB. However, schools of late-stage larvae of some herring and jack species can occur 

nearshore. Some herring and anchovy species, including species such as the thread herring 

(Opisthonema oglinum) and Anchoa spp., can form schools of elongate, translucent late-stage larvae in 

the water-column above NHB and IHB structure. 

Jacks are a diverse family with many representatives associating with NHB, IHB and OHB. Early life 

stages of many of the jacks, including Caranx spp., bigeye scad, and round scad, associate with drifting 

flotsam and Sargassum and utilize nearshore environments, including NHB, following strong easterly 

winds. Others, particularly Atlantic bumper, associate with large jellyfishes and are similarly transported 

from the open ocean into NHB areas. 

As mentioned above, most jacks species, including scads (Selar crumenophthalmus; Decapterus spp.), 

crevalles (Caranx spp.), amberjacks (Seriola spp.), Atlantic bumper, and rainbow runner (Elagatis 

bipinnulata), settle and associate with flotsam and Sargassum. Individuals from another group of jacks, 

consisting of Florida pompano (Trachinotus carolinus), permit (T. falcatus), and palometa (T. goodei), 

settle in very shallow coastal water (Fields, 1962) and will opportunistically shelter in tide pools formed by 

NHB, such as in Brevard County NHB (Continental Shelf Associates, Inc., 2005c). 
Section 2 5-13 

Organismal Assemblages of East Florida Shallow Hardbottom 

Baron et al. (2004) observed seven species from five genera of jacks. In their surveys of NHB and OHB, 

the bar jack (Caranx ruber) accounted for most of the observations and ranged from 3 to 25 cm TL. To 

the north, yellow jack (C. bartholomaei) was the most commonly observed carangid on NHB (Lindeman 

and Snyder, 1999). 

5.4.2.3 Juvenile and Adult Habitat Use 

Clupeid and engraulid records from all survey methods in the Fort Lauderdale and Jupiter study areas are 

absent or very low, respectively. Only Spanish sardine and unidentified engraulids were recorded from 

Jupiter NHB transects (scaled sardine was also found over open sand, near NHB). Based on roving diver 

surveys and on-water boat observations, the diversity and abundance of highly motile, surface-associated 

herring and anchovy species in association with NHB is underestimated by transect and point-count 

methods. Continental Shelf Associates, Inc. (1992) collected juvenile bigeye scad in mixed schools of 

juvenile clupeids associated with NHB in Jupiter and Palm Beach. 

Transect counts of jacks (carangids) at Fort Lauderdale sites found bar jack, yellow jack, and blue runner 

(Caranx crysos) at minimum sizes of 3, 7, and 7 cm TL, respectively. Mean lengths surveyed for these 

species were 8, 15, and 14 cm SL, respectively (Baron et al., 2004). Many of these individuals were 

juvenile stages, with bar jack in association with shallow hardbottom structure at post-settlement stages. 

Juveniles of these species were also recorded in the Jupiter surveys where divers observed 

post-settlement behavior in yellow jack at 3 cm. Schools of thousands of scombrids and bluefish are 

often fished in IHB but have not been censused. 

An exceptional example of a coastal pelagic schooling species in association with shallow hardbottom 

exists at the Peck’s Lake reef system in southern Martin County, where a semi-continuous gradient of 

worm-rock encrusted hardbottom extends from NHB through OHB depths and is used by a massive 

seasonal adult aggregation of Spanish mackerel (Scomberomorus maculatus). This aggregation has 

been highly predictable in the late fall through winter (October to late March) and heavily fished for well 

over a decade. There is little direct information on what the aggregating mechanism is; perhaps it is 

feeding. There is no evidence of spawning in the Peck’s Lake system, with spawning probably occurring 

later in spring and further offshore. The area is characterized by expansive NHB and IHB reefs, some 

with high (2 m) and convoluted relief, many with substantial P. lapidosa habitat structure, most of it the 

remaining framework of dead colonies. Unlike many NHB systems, hardbottom extends to approximately 

8 m depths offshore. 

Before the highly touted and exploited Spanish mackerel fishery at Peck's Lake reef, there was a massive 

seasonal aggregation of mackerel and bluefish inside the Lake Worth Lagoon, 45 km to the south. These 

fish entered Lake Worth usually during March and April, presumably on their way back north following the 

regular southerly migration triggered by the passage of winter cold fronts. This subpopulation was fished 

regularly by hundreds of small commercial hook-and-line fishermen, maintained consistency for years, 

and was considered a limitless resource. The aggregation began to diminish by the mid-1970’s with the 

growth of the commercial fishery, and also because of water quality degradation in the lagoon from 

increased dredge-and-fill activity in the lake and the expansion of the Port of Palm Beach (T. Twyford, 

West Palm Beach Fishing Club, West Palm Beach, FL, pers. comm.). When the fishery evolved beyond 

hook-and-line fishers, big net boats with roller rigs, spotter planes, and run-around gill nets heavily fished 

mackerel up and down the east Florida coast. The fishery begin to rebound through tighter gear 

restrictions and quota limitations, then the net ban in 1995 further boosted mackerel recovery. It appears 

that since the net ban, Spanish mackerel have been able to stay in the Pecks Lake area for long 

stretches with less disturbance from the current heavy hook-and-line fishing than from the run-around 

nets used before the net ban. Run-around nets also took other coastal pelagic species that associate 

with NHB, including ladyfish (Elops saurus) and jack crevalle (Caranx hippos). 

There are reports of a sizable aggregation of small Gulf kingfish (Menticirrhus littoralis) near Jap Rock in 

Boca Raton that occurs February/March; it is not as large as the Peck’s Lake aggregation and receives 

less fishing pressure (T. Gibson, Outdoor Life Magazine, pers. comm.). King mackerel (Scomberomorus 

cavalla) form similar aggregations at various sites further out on the shelf and shelf edge, as do the little 
Section 2 5-14 

Organismal Assemblages of East Florida Shallow Hardbottom 

tunny (Euthynnus alletteratus), but aggregations of neither species maintain as long as the Spanish 

mackerel do at Pecks Lake (D. Snyder, CSA International, Inc., pers. comm.). There are a few other 

predictable Spanish mackerel aggregations that also occur seasonally in Florida Bay and on Florida's 

southwest coast in the winter (T. Twyford, West Palm Beach Fishing Club, West Palm Beach, FL, pers. 

comm.). 

Each year in the fall, adult and subadult striped mullet (Mugil cephalus) mass migrate along the coast of 

southeastern Florida. Although not directly associating with NHB, the large schools of mullet pass near 

and over hardbottom areas during their migration. 

Adult bull sharks were observed at Coral Cove in northern Palm Beach County during July 2004 

(Continental Shelf Associates, Inc., 2005b). Large schools of blacktip and spinner sharks migrate along 

the east Florida coast during winter and early spring (D. Snyder, CSA International, Inc., pers. observ.). 

Groups of individuals from larger schools will associate with NHB temporarily (for a few days or weeks) 

during the migration. Large individuals of great hammerhead (Sphyrna mokkaran) are occasionally 

observed following the migrating blacktips (D. Snyder, CSA International, Inc., pers. observ.). 

5.4.3 Trophic Patterns 

Many coastal pelagic species are prominent piscivores, typically transient but capable of substantial 

localized predation in deeper NHB through OHB habitats. Prominent piscivorous species include the 

jacks, mackerels, cobia, and requiem sharks. Many piscivorous species also can feed on invertebrates. 

In Randall (1967), approximately half of the species of jacks fed on bottom macroinvertebrates, including 

shrimp, cephalopods, and crabs, while the gut contents of the other half of the species indicated they 

were piscivorous. The most abundant species in both the Jupiter and Fort Lauderdale surveys were the 

yellow jack and bar jack, species recorded to feed on invertebrates and fishes and only fishes, 

respectively. 

The piscivorous coastal pelagic subgroup includes some gamefish species that are preyed upon by 

seasonal aggregations of marine mammals (dolphins) that gather in coastal NHB through OHB 

environments in the fall and winter to feed on Spanish mackerel, bluefish, and cobia. These same 

species are also targeted by recreational anglers. The mackerel and bluefish are on the seasonal 

southern boundary of their fall migration to warmer water prior to late spring spawning off east Florida. 

Sharks are also routinely fished and sighted by anglers and surfers in deeper NHB and OHB areas; these 

species include nurse, blacktip, spinner, bull, lemon, sandbar (C. plumbeus), sharpnose (Rhizoprionodon 

terraenovae), and tiger sharks. 

Primary plantivorous coastal pelagic species include the Spanish sardine, round scad, bigeye scad, and 

thread herring (Opisthonema oglinum). In the northeast Caribbean, thread herring fed on copepods and 

also the planktonic stages of annelids and crustaceans (Randall, 1967). 

5.4.4 Latitudinal Distribution 

Observations of coastal pelagic species utilizing NHB in Fort Lauderdale (Baron et al., 2004) and Jupiter 

(Lindeman and Snyder, 1999) were limited to the jacks previously discussed. Differences in the numbers 

and kinds of coastal pelagic species between these two studies are difficult to explain because the visual 

sampling methods used do not adequately sample such highly migratory species. 

Some of the most prominent and seasonally periodic coastal pelagic species are warm temperate 

migrants from cooler, Carolinian waters of the South Atlantic Bight. Examples of seasonal transient 

schoolers that range from the shelf edge through NHB structure include bluefish (Pomatomus saltatrix) 

and Spanish mackerel. These species can probably be significant predators on OHB through deeper 

NHB habitats in the northern study area when large schools migrate off east-central Florida in fall months. 

However, these species typically do not go as far south as the NHB in Broward and Miami-Dade 

Counties. 
Section 2 5-15 

Organismal Assemblages of East Florida Shallow Hardbottom 

5.5 HERBIVORES AND OMNIVORES 

5.5.1 Focal Families and Species 

Many schooling herbivores common to Caribbean reefs occur in association with NHB and include the 

surgeonfishes (Acanthuridae), parrotfishes (Scaridae), porgies (Sparidae), and chubs (Kyphosidae); most 

of these families are represented largely by juvenile life stages, but also frequently occur as adults 

(Lindeman and Snyder, 1999; Baron et al., 2004). Non-schooling herbivores on NHB include the 

damselfishes (five species), filefishes (Monacanthidae), spiny puffers (Diodontidae), presumably all 

gobiids, and several blenniids (Randall, 1967). 

Herbivorous fishes may be defined as browsers on macroscopic algae and grazers on turfing algae 

(e.g., Choat et al., 2002; Floeter et al., 2005). In NHB environments, herbivores represented few species, 

but these can be locally abundant. Browsers are represented by the chubs (Kyphosus sectatrix and 

K. incisor) and adult porgies (Diplodus spp.). The chubs will also feed in surface waters on drifting 

Sargassum algae. Grazers are represented by surgeonfishes (Acanthurus spp.) and parrotfishes 

(primarily S. rubripinne). 

There can be substantial ontogenetic variation in feeding among some species. For example, the silver 

porgy (Diplodus argenteus), a very abundant sparid on NHB, is probably omnivorous like its closely 

related congener, the spottail pinfish, D. holbrooki. The latter species was found to eat both algae and 

macroinvertebrates by Hay and Sutherland (1988) in a study of rocky jetty-type structures in Murell’s Inlet, 

South Carolina. Carr and Adams (1972) documented that D. holbrooki fed on ectoparasites and were 

therefore considered to be cleaners. Juveniles of D. argenteus have been observed cleaning larger 

fishes in NHB environments (D. Snyder, CSA International, Inc., pers. observ.). 

5.5.2 Shelter Use 

5.5.2.1 Spawning Areas and Larval Transport 

Nearshore spawning in most motile and 

site-associated herbivores is not well known. There is 

considerable information from other locations and 

typically deeper, more coralline habitats for 

damselfishes. At least four damselfish species 

(sergeant major [Abudefduf saxatilis], cocoa 

damselfish [Stegastes variabilis], dusky damselfish 

[S. adustus], and beaugregory [S. leucostictus]) can 

spawn directly on NHB surface to build their egg nests 

(Figure 5.9). These species can clear algae from the 

substrata and lay adhesive demersal eggs directly 

onto the rock surface. 

It is difficult to summarize the use of NHB habitat for 

spawning in non-nesting species of herbivorous fishes 

since research information is very limited. Many 

species exhibit larger adult populations on deeper 

reefs, but the complexity is amplified in areas of the study region where NHB is not discrete, but is instead 

part of a continuous IHB through OHB habitat gradient. Other prominent taxa that can potentially spawn 

in association with NHB include porgies (Sparidae). 

Many other herbivores that use NHB are primarily represented by early life stages and typically do not 

spawn in association with NHB, though they may spawn in association with OHB. This includes some 

species of scarids and acanthurids. 

Figure 5.9. Example of nest of the sergeant 

major (Abudefduf saxatilis) on 

hardbottom, 20 m depth, Jupiter, 

Palm Beach County, Florida. 

Photo by D. Snyder.
Section 2 5-16 

Organismal Assemblages of East Florida Shallow Hardbottom 

5.5.2.2 Settlement and Early Associations with Shelter 

At least 20 species of herbivorous fishes settle on NHB. The more abundant of these include some of the 

most well known reef fishes of the Caribbean: sergeant major, cocoa damselfish, redfin and bucktooth 

parrotfishes (Sparisoma rubripinne and S. radians, in northern and southern sites, respectively), and 

doctorfish and ocean surgeonfish (Acanthurus chirurgus and A. bahianus, respectively). Seasonal 

recruitment pulses of these and other herbivores may simultaneously influence algal assemblage 

composition and provide a prey base for larger piscivores. 

Herbivorous species of damselfishes and surgeonfishes have displayed substantial temporal variation in 

larval recruitment studies in other regions, and these patterns have been used to argue that coral reef 

fishes represent non-equilibrium systems (Sale, 1991). Notably, these studies are typically from deeper 

and more coralline habitats. It is logical to also treat NHB habitats as non-equilibrium systems that are 

highly mediated by disturbances that impact algal and herbivore abundance in unstudied manners. 

Most of species of scarids (parrotfishes) settle on NHB or IHB. Baron et al. (2004) recorded stoplight 

(Sparisoma viride) and bucktooth parrotfishes as small as 2 cm and six other species including Scarus

spp. and bluelip parrotfish (Cryptotomus roseus) as small as 3 cm, strongly suggesting that these 

individuals settled on NHB or OHB in their study area. Lindeman and Snyder (1999) observed newly 

settled silver porgy that settled predictably in relatively high numbers during late winter and early spring. 

Damselfishes and chubs do not appear to have particularly distinct settlement seasons but were 

sporadically observed as newly settled individuals. 

5.5.2.3 Juvenile and Adult Habitat 

Shelter use characteristics of herbivores largely involve daytime site associations and social behaviors 

that can be considered in terms of either site-associated species or wide-ranging, often schooling 

species. The primary site-associated herbivores include at least two different groups: the damselfishes, 

which can enforce relatively sizeable territories to maintain algal feeding sites, and species of gobiids and 

blenniids. The feeding of the latter group is less known than in damselfishes, but is assumed to be 

associated with short-distance benthic browsing movements. More motile and gregarious species also 

feed on NHB macroalgae, including surgeonfishes, parrotfishes, and chubs. These species can form 

sizable schools that effectively remove macroalgal cover on deeper reefs. 

The numerical contribution of herbivores to the assemblages characterized by Baron et al. (2004) in 

Fort Lauderdale and Lindeman and Snyder (1999) in Jupiter differed in terms of overall diversity and 

abundance of dominant taxa. Censuses conducted in Jupiter (Lindeman and Snyder, 1999) revealed that 

26% of the individuals observed were herbivores, numerically dominated by (in rank order of abundance) 

silver porgy, cocoa damselfish, ocean surgeon, doctorfish, redfin parrotfish, redtail parrotfish 

(S. chrysopterum), and juvenile parrotfish (Sparisoma spp.). The diversity of herbivores, especially the 

parrotfishes, was higher in the southern area studied by Baron et al. (2004). They observed 17 herbivore 

species, including damselfishes, porgies, surgeonfishes, parrotfishes, and chubs; this group was 

dominated numerically by bucktooth parrotfish. Parrotfishes represented 2% of the individual 

abundances recorded in transects by Baron et al. (2004). Lindeman and Snyder (1999) found higher 

abundances of chubs, porgies, and cocoa damselfishes in Jupiter. 

5.5.3 Trophic Patterns 

Early life stages of schooling herbivores frequently occur in association with NHB and include the 

surgeonfishes (Acanthuridae), parrotfishes (Scaridae), chubs (Kyphosidae), and porgies (Sparidae) 

(Figure 5.10). Available evidence suggests that all the species of the three former families are almost 

wholly herbivorous (Randall, 1967; Sierra, 2001), but the porgies are not. The seasonally abundant silver 

porgy, D. argenteus, fed on algae, gastropods, and crabs in the northeast Caribbean and is omnivorous 

(Randall, 1967). Non-schooling herbivores that are residents on NHB include the damselfishes 

(five species), some species of filefishes, one species of spiny puffer, several blenniids, and perhaps all 

of the gobiids (Randall, 1967). 
Section 2 5-17 

Organismal Assemblages of East Florida Shallow Hardbottom 

There are few or no feeding 

studies on these families from 

east Florida hardbottom habitats. 

In the northeast Caribbean, all 

species of Stegastes are 

omnivorous, with some annelids 

and crustaceans as well as algae 

seen in their gut contents. 

Beaugregory and threespot 

damselfish (Stegastes planifrons) 

were found to have the most algae 

in their diet per volume (Randall, 

1967). Sergeant major also is 

omnivorous, feeding on algae, 

anthozoans, and crustaceans. 

Based on Randall (1967), 

omnivores include 1) species of 

filefishes that feed on hydrozoans, 

algae, gorgonians, and 

zooantharians, and 2) the 

sharpnose puffer, Canthigaster 

rostrata, which feeds on plants, sponges, echinoids, and molluscs. 

Hay and Sutherland (1988) state that in warm temperate, rocky nearshore habitats, well-lit areas are 

algae-dominated and shaded areas are the realm of invertebrates. In east Florida, we cannot falsify this 

since well-lit open areas of NHB can be dominated by algae (though P. lapidosa encrustations can also 

dominate in some locations). This is presumably one reason that from a distance, the ichthyofauna on 

the well-lit crests of shallow hardbottom can appear to be dominated by herbivores (chubs, porgies, 

damselfishes), while many invertivores are located near-bottom, associated with microhabitats away from 

the most well-lit and open areas. 

5.5.4 Developmental Patterns and Cross-shelf Habitat Use 

Resident herbivores such as damselfishes and cryptic hardbottom species do not engage in ontogenetic 

habitat shifts. Some schooling species (surgeonfishes, parrotfishes, or chubs) represented largely by 

juveniles on NHB may migrate away from NHB to deeper structural habitats with maturation, but definitive 

information is unavailable. The ability to successfully make such migrations will correlate in complex 

manners with the degree to which cross-shelf habitat corridors are available and also drivers associated 

with potential ontogenetic shifts in food needs. 

5.5.5 Latitudinal Distribution 

There are complex similarities and differences in the presence and abundance of herbivores between the 

two sites for which quantitative data are available. Cocoa damselfish was the most abundant 

pomacentrid in both studies, however, the closely-related beaugregory was much more abundant at the 

southern site. Sergeant major were abundant in both areas (Lindeman and Snyder, 1999; Baron et al., 

2004). 

The silver porgy (represented primarily by juveniles, which are not herbivorous) was the second-most 

abundant species at the northern Jupiter sites but was not a significant component of the Fort Lauderdale 

assemblage (Lindeman and Snyder, 1999; Baron et al., 2004). More parrotfish species were found in the 

Fort Lauderdale study, probably due to the wider depth distributions of the surveys. Floeter et al. (2005) 

documented a negative correlation between diversity and density of herbivorous fishes with latitude, thus 

the low numbers of herbivorous species observed on NHB may reflect a larger-scale pattern. 

Figure 5.10. Early juvenile stages of reef herbivores including 

surgeonfishes, damselfishes, and parrotfishes on 

nearshore hardbottom, 3 m depth, Palm Beach, 

Palm Beach County, Florida. Photo by D. Snyder.
Section 2 5-18 

Organismal Assemblages of East Florida Shallow Hardbottom 

5.6 SUMMARY 

5.6.1 Species Richness 

During the present study, it was possible to revisit and expand upon the most recent fish species lists for 

NHB of mainland east Florida. Based on new and old information, including the work of Baron et al. 

(2004) and Herrema (1974), over 250 species of fishes are now recorded from NHB (Appendix C, 

Table C.1). This list documents species records and spatial and temporal attributes for all fish species 

recorded from NHB. Similarities with many shallow coral reef assemblages are present (Banks et al., 

2008). Differences between NHB fish assemblages and hermatypic reef assemblages in OHB depth 

ranges can involve the higher relative proportion of early life history stages on NHB structures, a 

juvenile-dominated assemblage (Lindeman and Snyder, 1999; Baron et al., 2004). 

The current working number of species is 257 (Appendix C, Table C.1) from 68 families. The previous 

list of fish species recorded from 0 to 4 m NHB totaled 192 species (Lindeman, 1997a). Primary additions 

to that estimate derive from the Baron et al. (2004) study in the southern project area in depths to 7 m, 

which added several dozen species to the list of records, many of which are uncommon or not found in 

0 to 4 m NHB in the northern project region. This pattern suggests that there is a southern NHB 

ichthyofauna and a less diverse northern fauna, particularly north of the deflection of the Gulf Stream 

offshore. We also characterized fundamental ecological attributes of these species (Appendix C,

Table C.1). Species assessments of trophic patterns, residency, size classes, life stages, and the 

components within from this table (e.g., higher percentages of piscivores and planktivores compared to 

herbivores) are ongoing. 

5.6.2 Spawning 

A wide range of spawning patterns is seen among the fishes of east Florida shallow hardbottom. Nesting 

is undertaken in differing manners by species of fishes from quite different families. Damselfishes, locally 

abundant herbivores, can influence subtidal fish distributions on 1-m scales by their aggressive

territoriality. Some of the species recorded from NHB build and protect nests in which they grow their 

young, sometimes at monthly intervals (Thresher, 1984). Several very different families, including the 

site-attached labrisomids, blennies, and gobies (potentially important predators of small fishes and 

invertebrates), also build nests in cryptic manners on NHB structure, often laterally or upside-down on 

inverted microhabitats. Though understudied, we estimate that over 30 fish species spawn in association 

with NHB. Most motile species do not spawn on NHB, though there probably are exceptions. In addition, 

some coastal pelagics spawn on or near NHB (Continental Shelf Associates, Inc., 1992). 

5.6.3 Settlement and Early Associations with Habitat 

Directed studies of fish settlement in NHB through OHB depths are lacking in the mainland east Florida 

study region. Patterns of settlement can be inferred from the larger, assemblage studies. In some 

species of grunts, there is evidence of potentially substantial variation in larval recruitment among years 

(Lindeman and Snyder, 1999; Jordan et al., 2004; Continental Shelf Associates, 2006; Coastal 

Eco-Group Inc., 2008). However, estimates of temporal patterns in fish settlement in the region (from 

seasonal to interannual) are currently based on very limited information. 

Information on the use of the water column by settlement-stage larvae is lacking. Larval occurrence of 

some taxa in surf zone areas has been examined in other areas but not in this study region (Ruple, 1984; 

Jahn and Lavenberg, 1986). Low or high relief hardbottom could be a visual cue for settling larvae, but 

other sensory systems, particularly hearing and olfaction, can also be important (Kingsford et al., 2002). 

Larval fishes, including many clupeids, engraulids, and atherinids (Sponaugle et al., 2003), can form 

assemblages over shallow coral reefs but are unstudied over NHB or IHB. Sponaugle et al. (2003) found 

that shallow, near-reef larval fish assemblages differed from offshore collections. A significant aspect of 

this was the higher nearshore frequency of haemulid larvae. 
Section 2 5-19 

Organismal Assemblages of East Florida Shallow Hardbottom 

Are nearshore fish larvae spending substantial time in the water column near NHB before settling there? 

This is possible for some taxa (e.g., clupeioids) but cannot be confirmed without more empirical support. 

For many species in South Florida, transport of larvae to NHB or OHB settlement areas may occur via 

cross-shelf oceanographic transport mechanisms resulting from diverse meso-scale phenomena in south 

and east Florida (Lee et al., 1995; Limouzy-Paris et al., 1997). 

5.6.4 Developmental Patterns and Cross-shelf Habitat Use 

Substantial stretches of NHB exist in portions of Broward, Palm Beach, Martin, St. Lucie, Indian River, 

and Brevard Counties (26°20' to 27°15' N) and elsewhere (Perkins et al., 1997; Table 2.1, present study). 

Although these areas are typically separated by kilometers of sand, there are some areas where there is 

a semi-continuous hardbottom gradient from the beach into depths exceeding 6 m. Offshore, many 

mid-shelf areas (5- to 20-m depths) are also dominated by expanses of sand, despite the variable 

occurrence of several mid-shelf reefs. Therefore, in many shallow areas, there are no natural habitats in 

the same or adjacent nearshore areas that can support equivalent abundances of early life stages of 

fishes and the invertebrates they largely feed on. Absences of nursery structure can logically result in 

increased predation and lowered growth. In newly settled and juvenile stages, such conditions could 

foster demographic bottlenecks that ultimately result in lowered local population sizes (Jones, 1991). 

Empirical information is highly limited on the amount of connectivity between shallow patches of 

hardbottom and deeper reefs for fishes and invertebrates of east Florida. The limited available 

information suggests that diverse fish species that can ontogenetically migrate into deeper water use 

NHB during early life stages (Gilmore et al., 1981; Vare, 1991; Lindeman and Snyder, 1999; Baron et al., 

2004). If the NHB in use is close enough to IHB or OHB, some of these species, depending on many 

changing biophysical drivers (e.g., predation, growth, larval recruitment) could migrate ontogenetically 

into deeper habitats. Answers to the question, “How close is close enough?” will vary by species, site, 

and time. For fishes, tagging studies would be useful using species on highly segregated, NHB-depth 

reefs in comparison to fishes tagged on NHB structure that is not restricted to depths <4 m but extends as 

emergent structure into depths of >6 m. 

NHB can be not only spatially patchy for shallow organisms, it can also be temporally patchy due to 

natural burial events. Natural burial events are typically associated with rapid sediment redistributions in 

association with high wind and/or wave events. These events and partial or complete NHB burial are 

more common in the winter. In the absence of some storm-induced burial of subpatches or patches of 

habitat, NHB can be subject to very high wave energy and substantial scouring of surfaces, especially in 

the intertidal and immediate subtidal. Many organisms assumedly would associate with shelter as 

possible during these events, perhaps moving into the deepest NHB or IHB or nearby sand area available 

and returning post-event, if feasible. Information on organismal responses to high wind and wave events 

is needed and will assist in discerning larger patterns of cross-shelf distribution. 

Are there substantive reasons to not consider the fishes associated with NHB a reef fish assemblage? A 

useful discussion on this topic in the literature occurred regarding benchline criteria for both taxonomic 

and ecological definitions of “what reef fishes are” (Bellwood and Sorbini, 1996; Bellwood, 1998; 

Robertson, 1998a,b). Bellwood (1998) concluded that reef fish assemblages ultimately occur in 

association with too many non-coral reef structures and recommended instead the term fishes on coral 

reefs. Robertson identified various challenges to a unifying taxonomic structure. He defined reef fishes 

as “species that live on consolidated substrata that form coral and inorganic reefs.” He described 

inorganic reefs, including rocky shores, as follows: “... the inorganic bedrock provides large and small 

physical structure that fishes use as habitat features...”. In addition, all of the 10 consensus families of 

reef fishes in Bellwood (1998) are represented in association with NHB of east Florida. Earlier sections 

detail the long-standing geological literature on the reef attributes of NHB (e.g., Chapters 1 and 2). 

These lines of evidence suggest there are few, if any, reasons to conclude that the fishes of NHB of 

mainland east Florida do not constitute a reef fish assemblage. 
Section 2 5-20 

Organismal Assemblages of East Florida Shallow Hardbottom 

5.6.5 Trophic Patterns and Functional Groups 

There are no feeding studies of fishes from shallow hardbottom habitats in the project region. The most 

detailed examination of food habits of Caribbean reef and coastal pelagic fishes remains Randall (1967). 

His study areas were in the U.S. Virgin Islands and Puerto Rico. In the last decade, a series of detailed 

studies of feeding and habitat use of grunts and snappers has occurred in Curacao and other Caribbean 

islands (e.g, Nagelkerken et al., 2000; Dorenbosch et al., 2004). The older and newer trophic studies are 

largely from insular systems with potential differences in seascapes and landscapes in comparison to 

continental, mainland shelf areas such as east Florida (Robins, 1991). Based largely on studies from 

insular areas, the grunts and margates of the genera Haemulon and Anisotremus show a wide array of 

invertivory. Some species are planktivorous as adults (H. striatum), others are very specialized for 

feeding on hard-shelled invertebrates (H. macrostomum), while the majority of species are relatively 

generalized feeders on both soft- and hard-invertebrate prey, often from either softbottom or hardbottom 

environments (Randall, 1967; Sierra, 2001). 

Most NHB species are demersal invertivores; however, assemblages also include important demersal 

piscivores and planktivores. Over 50 species of NHB fishes consume invertebrates as a primary dietary 

component. Eleven species of wrasse exihibit a pattern similar to grunts and other families: invertebrates 

from many diverse taxa are consumed by differing congeners with low to high levels of specialization 

within the genus. For example, many wrasse species (genus Halichoeres) feed on “shelled invertebrates” 

in the northeast Caribbean, though specific dietary compositions can vary (Randall, 1967; Randall’s data 

tabulated by taxa in Sierra, 2001). The dominant labrid in both the Jupiter and Fort Lauderdale studies 

was the slippery dick (H. bivittatus). This species feeds on crabs, urchins, annelids, and gastropods 

(Randall, 1967) and is an opportunistic feeder. Other labrids, such as the clown wrasse, H. maculipinna, 

primarily feed on soft annelids (Randall, 1967). As mentioned elsewhere, these comparative patterns are 

largely predictive without empirical feeding information from the project region. 

Although the NHB assemblage is juvenile-dominated, a well-developed food web exists, with all of the 

trophic categories of Bellwood’s (1998) “consensus reef fish families” represented, including diverse 

herbivores and piscivores on juvenile fishes. Following from Randall (1967) and Bohnsack and Harper 

(1988), we characterized fish species according 

to the following trophic categories: herbivore, 

planktivore, detritivore, invertivore, and piscivore. 

This categorization scheme aids assessment of 

potential functional groups to better understand 

trophic relationships among NHB, IHB, and OHB 

assemblages. When the full spectrum of 

ontogenetic dietary shifts is considered for most 

species, many are omnivorous (e.g., planktivory 

to invertivory to piscivory from species as diverse 

as labrisomids and lutjanids). The degree of 

feeding variation within individual genera can be 

high, and it is very important to recognize that 

feeding patterns can vary geographically based 

not only on variable within-species behavior but 

based also on geography-dependent food 

availability (Turingan et al., 1995) and 

inter-annual variations in food availability (Sierra, 

2001). 

The most prevalent trophic component among 

species was invertebrate feeding (Figure 5.11). 

Invertivory was four times more common among 

species than herbivory or planktivory. Randall categorized the diversity of invertivores according to 

1) sessile animal feeders (e.g., sergeant major feeding on Zooanthus), 2) “shelled” invertivores (e.g., most 

Halichoeres species of wrasses), and 3) generalized invertebrate predators. The latter group was by far 

Figure 5.11. Relative adult trophic composition 

as measured by numbers of species 

per functional grouping. 
Section 2 5-21 

Organismal Assemblages of East Florida Shallow Hardbottom 

the biggest and included eels, squirrelfishes, snappers, species of basslets (Serrranus), some drums 

(sciaenids), goatfishes (mullids), pompanos (Trachinotus spp.), clown wrasse (H. maculipinna), and some 

labrisomids, including the hairy blenny. Specialized invertebrate feeding can occur as well, as in the 

preference for echinoids in older black margates. Early juvenile stages of porkfish, Anisotremus

virginicus, and porgies, Diplodus spp., can serve as cleaners of invertebrate ectoparasites. 

Piscivory ranked almost twice as high as herbivory and planktivory (Figure 5.11) but is inflated by the 

presence on the list of many piscivorous pelagic species that uncommonly associate with NHB. 

Prominent piscivores include lizardfshes (synodontids), jacks, snappers, muraenid and ophichthid eels 

(Gymnothorax and Ophichthus spp.), mackerels, and sharks. Many diverse piscivores feed on 

invertebrates as well (e.g., tarpon, nurse sharks) (Randall, 1967). 

Herbivores were represented by juvenile stages of many common schooling herbivore species of 

Caribbean reefs and included the surgeonfishes (three species of Acanthurus), parrotfishes (especially 

redfish parrotfish), silver porgy (Diplodus), and chubs (Kyphosus incisor and K. sectatrix) (Appendix C, 

Table C.1). Less mobile herbivores include the damselfishes (five species). Omnivorous species of 

filefishes and puffers, all gobiids, and several blennioids are also herbivorous (Randall, 1967). 

Planktivorous fishes of shallow hardbottom are diverse and include over 10 species of herring and 

anchovies (clupeids and engraulids), scads (Decapterus spp. and Selar crumenophthalmus), and a 

variety of demersal fishes including species of apogonids, blackbar soldierfish (Myripristis jacobus), and 

small demersal schools of copper sweeper (Pempheris schomburgkii). Some locally abundant NHB 

species, including thread herring (Opisthonema oglinum) and copper sweeper, were found by Randall 

(1967) to feed heavily on polychaete larvae (23% of total volume, 23 specimens and 27%, 19 specimens, 

respectively) in the northeast insular Caribbean. Substantial planktivory also occured in species of drums 

(Pareques spp. and Odontoscion dentex), sergeant major, bluehead wrasse, and juvenile stages of 

haemulids and carangids (Randall, 1967). The species-specific and collective impacts of these 

planktivores upon the seasonal dynamics of worm reef formation by P. lapidosa larvae deserve study. 

5.6.6 Latitudinal Variations in Shallow Hardbottom Fish Assemblages 

Some differences among ichthyofaunal assemblages are present between the southern and northern 

areas of mainland southeast and east-central Florida in terms of the most abundant species and habitat 

structure (e.g., earlier examples in present chapter; Appendix C, Table C.1). Factors influencing these 

potential differences include 1) differences in the quantity or structure of shallow hardbottom of southeast 

Florida compared to east central Florida and 2) major, long term differences between southern and 

northern sites in presence/absence or abundance of locally prominent species. The available NHB areal 

data shows higher cover in some areas compared to others (Table 2.1). In the second case, there are 

not easy numeric or image-based answers. In terms of assemblages, the differences could be based on 

true differences in fish assemblages between southeast Florida and east-central Florida or sampling 

artifacts, differing physical regimes, larval recruitment variation, post-settlement growth and mortality 

variation, or combinations of these factors. 

There are significant differences in abundance between three of the most abundant species in the Jupiter 

area in comparison to the Fort Lauderdale area. Only one black margate out of >72,000 individuals was 

recorded in Fort Lauderdale (Baron et al., 2004), while 636 black margates out of 10,491 individuals were 

censused in Lindeman and Snyder (1999). Also, 43 sailors choice were recorded in Baron et al. (2004) 

with mean lengths above 15 cm, while 1,326 H. parra individuals were recorded in Lindeman and Snyder 

(1999), primarily early juvenile stages below 5 cm. The hairy blenny was the most abundant cryptic 

hardbottom fish species in the Jupiter surveys, with 806 individuals recorded in transect surveys, 8% of 

the overall individual total (Lindeman and Snyder, 1999). Though the hairy blenny was recorded in 

qualitative roving diver surveys, none were recorded in transect and point-counts in Fort Lauderdale 

(Baron et al., 2004). 
Section 2 5-22 

Organismal Assemblages of East Florida Shallow Hardbottom 

Potential explanations for such marked difference in dominant species within the 100-km north-south 

distance between these studies may include temporal recruitment variations, NHB structural differences 

between survey areas, species reaching their biogeographic distributional limits between these sites, and 

other factors. The Fort Lauderdale study occurred during several summer months in 2001, while the 

Jupiter study occurred between 1994 and 1996. Recruitment in haemulids can vary across temporal 

scales (McFarland et al., 1985; Jordan et al., 2004), and the possibility that the patterns reflect differences 

in interannual larval recruitment in the 6 years between these survey snapshots cannot be excluded. 

NHB structural differences probably exist between these areas as described above, but many of the more 

prominent species appear to be relatively opportunistic in their use of differing NHB and IHB microhabitat 

structure. At least three of the prominent species (black margate, sailors choice, and hairy blenny) with 

apparent major distributional differences here are listed by Gilmore and Hastings (1983) as species 

whose Florida distributions are restricted to east Florida, not occurring in the Gulf of Mexico as congeners 

do. More research is required to parse the multiple factors and examine the above alternatives to better 

understand these and other north-to-south differences in fish assemblage pattern. 
Section 2 6-1 

Organismal Assemblages of East Florida Shallow Hardbottom 

Chapter 6

Marine Turtles

6.1 INTRODUCTION 

Much of what is known about marine turtles in Florida is from their nesting habits on sandy beaches. 

More recently, acknowledgment has been made regarding the importance of NHB as important habitat for 

juvenile and subadult species of marine turtles. Green (Chelonia mydas), loggerhead (Caretta caretta), 

and hawksbill (Eretmochelys imbricata) turtles utilize NHB as developmental habitat in waters around 

Florida. These species have fundamental developmental requirements that are supplied by nearshore 

hardbottom (e.g., food, shelter, predator avoidance). Data from mark and recapture studies indicate that 

turtles may revisit or take up long-term residency on nearshore reefs (Holloway-Adkins, in prep.). Impacts 

of coastal development on marine turtles include pollution, dredge and fill projects, shoreline hardening, 

and commercial and recreational fishing activities. This document explores the use of nearshore waters 

by adult marine turtles and their hatchlings; however, particular focus is directed towards the use of NHB 

as developmental habitat for juvenile stages of marine turtles. 

6.1.1 Adult Marine Turtles 

Nearshore waters act as important corridors during the life history stage of nesting adult females and 

emerging marine turtle hatchlings. The terrestrial encounter with sandy beaches for nesting represents 

but a brief moment in the life history of all marine turtles; however, due to the ease of accessibility, it has 

been the most studied. Populations of adult female marine turtles use nearshore waters to access 

adjacent beaches during turtle nesting seasons. Depending on the species, nesting in Florida can begin 

as early as February for leatherback turtles (Dermochelys coriacea) and end as late as November for 

green turtles. 

NHB exists adjacent to many of the prime nesting beaches on the east Florida coast. Rocks are exposed 

during low tide events on some of these beaches; however, no historical evidence exists indicating NHB 

rock impedes marine turtle nesting in these areas. Turtles have been observed climbing over exposed 

rocks while entering and exiting the surf during nesting events (Figure 6.1). 

Figure 6.1. Adult female green turtles (in foreground and distance) crossing the nearshore hardbottom 

structure after nesting in Coral Cove Park, Palm Beach County, Florida. Photo by D. Snyder. 
Section 2 6-2 

Organismal Assemblages of East Florida Shallow Hardbottom 

6.1.2 Hatchling Marine Turtles 

A generalized conceptual model of ontogenetic habitat stages for marine turtles was constructed by 

Musick and Limpus (1997) that describes the functional role of oceanic and neritic habitats in the life 

history of marine turtles (Figure 6.2). The life cycle for most marine turtle species begins on a subtropical 

or tropical beach. Marine turtles hatch from eggs inside clutches of eggs laid and incubated in warm 

summer sands. Newly hatched turtles emerge from the nest and make their way across the sand and 

into the ocean. Turtles that hatch from Florida’s Atlantic beaches are believed to spend the next 3 to 

11 years (highly variable among species) in the pelagic environment, riding eddies and other circulatory 

systems within the North Atlantic Gyre (Figure 6.3). Juvenile green turtles enter neritic habitats at around 

20 to 25 cm straight carapace length (SCL). Data from age-at-size models indicate their age to be 

approximately 3 to 5 years (Witherington et al., 2006a). Loggerheads are rarely encountered in 

nearshore environments until approximately 45 cm SCL. Their time at large is estimated to be between 

6.5 and 11.5 years (Bjorndal et al., 2000). 

Figure 6.2. Conceptual model of ontogenetic habitat stages in marine turtles (From: Musick and Limpus, 

1997). 
Section 2 6-3 

Organismal Assemblages of East Florida Shallow Hardbottom 

Figure 6.3. North Atlantic Gyre with potential oceanic transport routes for juvenile marine turtles 

(From: Musick and Limpus, 1997). 

More recently, adjacent hardbottom habitats have been recognized as juvenile developmental habitat 

(Wershoven and Wershoven, 1992a; Bresette et al., 1998; Ehrhart et al., 2001; Holloway-Adkins et al., 

2002; Jones et al., 2004; Makowski et al., 2006b; Garrido, 2007). Few data exist on populations of 

marine turtles utilizing NHB as developmental habitat, and not enough data are available to clearly 

establish long-term patterns in the distribution and abundance of marine turtles associated with NHB on 

the east Florida coast. Data herein are predominantly from non-peer reviewed articles, reports, and 

personal communications and observations, though more information is beginning to enter the 

peer-review literature. Table 6.1 lists marine turtle species and their presence in or around NHB and 

indicates areas that are still in question. 

6.1.3 Predators 

Many predators prey upon hatchlings even before they leave the nest. Frequently, ghost crabs, beetles, 

worms, and other invertebrates invade the nest before hatchlings emerge to the surface of the sand. 

Raccoons, coyotes, and vultures are capable of destroying nests and consuming eggs and hatchlings. 

If a hatchling survives exiting the nest, reaches the surface, and makes its way to the sea, a ghost crab, 

sea bird, or a raccoon may consume it. Once in the water, the long journey out to meet the Florida 

Current is wrought with more predators that are capable of making a meal of a small hatchling turtle. 

Interestingly, Whelan and Wyneken (2007) found no significant differences between predation levels of 

hatchlings migrating from nearshore waters over either sand or hardbottom nearshore waters; however, 

higher hatchling predation levels were documented for animals that migrated over the more predator-rich 

hardbottom habitat. As marine turtles mature, their largest threat comes in the form of discarded 

anthropogenic debris that turtles may ingest (Bjorndal et al., 1994) or become entangled in and drown. 

Impacts from commercial fishing activities (e.g., drowning in shrimp trawler nets, long-line fishing gear, 

and gill nets) pose the greatest threats to loggerhead and Kemp’s ridley (Lepidochelys kempii) turtles. 
Section 2 6-4 

Organismal Assemblages of East Florida Shallow Hardbottom Table 6.1. Role (functions) of nearshore hardbottom (NHB) (and reference to offshore hardbottom [OHB]) in marine turtle 

Species Life Stage NHB OHB Diet Shelter Corrid

hatchling no no n/a no yes 

post no no omnivorous no yes 

juvenile yes yes omnivorous - herbivorous yes yes 

subadult no no herbivorous no possib

Green turtle (Chelonia mydas) adult no no herbivorous OHB – nesting? yes 

hatchling no no n/a no yes 

post no no omnivorous no yes 

juvenile yes yes carnivorous possible probably 

subadult yes yes carnivorous possible possib

Loggerhead turtle (Caretta caretta) adult yes yes carnivorous OHB – nesting yes 

hatchling no no n/a no possib

post no no omnivorous no no 

juvenile yes yes spongivorous yes possib

subadult yes yes spongivorous yes possib

Hawksbill turtle (Eretmochelys imbricata) 

adult no no spongivorous OHB – nesting? possib

hatchling no no n/a no yes 

post no no jellyfishes no no 

juvenile no no jellyfishes no no 

subadult no no jellyfishes no no 

Leatherback turtle (Dermochelys coriacea) 

adult yes yes jellyfishes no possib

n/a = Not available. 
Section 2 6-5 

Organismal Assemblages of East Florida Shallow Hardbottom 

6.1.4 Water Depth 

Visual transect surveys to determine turtle density distribution along NHB were conducted by Inwater 

Research Group, Inc. (2005) in St. Lucie and Martin Counties. Transects were surveyed for turtles 

inshore (200 m from shore, 3 m water depth) and offshore (400 m from shore, 4 m water depth) waters. 

Inwater Research Group, Inc. found that loggerhead turtles were similarly distributed across inshore and 

offshore surveys. However, green turtle densities were twice as high inshore as offshore. It is possible 

that loggerheads select deeper reef and sand bottom habitats to forage and rest. Their food resources 

are available in both habitat types. Visual transect surveys conducted in Brevard (Holloway-Adkins and 

Provancha, 2005) and Indian River (Inwater Research Group, Inc., 2004) Counties indicate the density of 

juvenile green turtles is higher in shallow nearshore waters. High catch-per-unit-effort (CPUE) rates are 

associated with the shallower reefs in 1.5 to 2.5 m water depths, closest to shore in Brevard and Indian 

River Counties (unpublished data). 

6.1.5 Seasonality 

Transect survey data indicate that marine turtles are present year-round at all of the NHB study sites 

(Inwater Research Group, Inc., 2004, 2005; Holloway-Adkins, 2005). Seasonal data from tangle net 

capture methods are normally restricted to summer months when ocean conditions are relatively calm. 

Tangle nets, however, are used to capture turtles entrained into the intake canals at the St. Lucie Power 

Plant in St. Lucie County. Seasonal data from the plant show a high number of very small size-class 

juvenile green turtles (potential new ocean recruits) are present each winter (Bresette et al., 1998). 

6.2 SPECIES OVERVIEWS 

Five of the seven circumglobally distributed marine turtles species have been observed in or adjacent to 

NHB of mainland east Florida. Largely depending on the time of year or latitude of observation, the most 

frequently observed turtles commonly are green, loggerhead, and hawksbill turtles. In deep offshore 

waters (>20 m), fishermen have reported frequent sightings of loggerhead turtles and occasional 

sightings of leatherback turtles. 

A small number of studies have investigated marine turtles in nearshore waters on the east Florida coast 

(Figure 6.4, Table 6.2). Two relatively long-term studies (i.e., 19+ years) have been conducted on 

marine turtles associated with NHB on the east Florida coast. Ehrhart et al. (2003) has researched 

marine turtles on sabellariid worm reefs near Sebastian Inlet in Indian River County, in summer, since 

1989. A long-term study of marine turtles entrained from offshore cool-water intake pipes at the St. Lucie 

Power Plant in St. Lucie County has been documented since 1976 (Bresette et al., 1998; Ecological 

Associates, Inc., 2000). 

Shorter or more recent studies have been conducted in Brevard County (Holloway-Adkins and 

Provancha, 2005), Palm Beach County (Makowski et al., 2006b) (L. Wood, Palm Beach Zoo, unpublished 

data), and Broward County (Wershoven and Wershoven, 1992a). Additional studies utilizing passive 

observation techniques to describe and/or assess marine turtle populations over NHB have been used in 

Indian River County (Inwater Research Group, Inc., 2004), St. Lucie and Martin Counties (Inwater 

Research Group, Inc., 2005), Palm Beach County (Jones et al., 2004; Garrido, 2007) and Broward 

County (Makowski et al., 2006a). Wood’s (2006) study is the first and only study to focus on hawksbill 

turtles on the east coast of Florida. 

There are few studies that utilize tangle net capture methods in NHB, but this method standardizes CPUE 

so comparisons among studies can be made. CPUE is based on the length of the net or nets used, the 

amount of time the net is in the water, and the number of turtles captured during that time. CPUE data 

indicate higher capture numbers per effort in Indian River County (Table 6.3). 
Section 2 6-6 

Organismal Assemblages of East Florida Shallow Hardbottom 

Figure 6.4. General locations of study areas for marine turtles in coastal areas superimposed over 

distribution of reef-building sabellariid worms along mainland east Florida (Modified 

from: Zale and Merrifield, 1989). 

Mid-Reach Nearshore Hardbottom Turtle Study – 

Brevard County 

Sabellariid Worm Reef Turtle Study – Indian 

River County 

St. Lucie Power Plant Water Flow 

Intake from Nearshore Reef Turtle 

Study – St. Lucie County 

Nearshore Reef Green 

Turtle Study – Palm 

Beach County 

Nearshore Reef Hawksbill 

Turtle Study – Palm Beach 

County 

Nearshore Reef Turtle 

Study – Broward County 
Section 2 6-7 

Organismal Assemblages of East Florida Shallow Hardbottom 

Table 6.2. Marine turtle mark and recapture population studies conducted over nearshore hardbottom. 

Marine Turtle Study Location 

(County) Capture Method Time of Year Duration Permit Holder (Current) Species 

Mid-Reach Brevard Tangle nets Mostly summer 2004 to Present Holloway-Adkins Cm, Cc 

Indian River County Indian River Tangle nets Summer 1989 to Present Ehrhart Cm, Cc, Ei 

St. Lucie Power Plant St. Lucie Entrainment 

and nets Year-round 1976 to Present Bresette Cm, Cc, Ei, Dc, Lk 

Little Breakers Reef Palm Beach Night SCUBA 

diving Summer and Fall 2003 to 2004 Wyneken Cm 

Breakers Reef Palm Beach SCUBA diving Year-round 2004 to Present Wood Ei 

Broward County Broward SCUBA diving Year-round 1987 Wershoven Cm, Cc, Ei 

Cm = Chelonia mydas; Cc = Caretta caretta; Ei = Eretmochelys imbricata; Dc = Dermochelys coriacea; Lk = Lepidochelys kempii. 

Table 6.3. Catch-per-unit-effort (CPUE) for marine turtle population studies conducted with tangle net 

capture methods adjacent to nearshore hardbottom habitats on the east coast of Florida. 

Location CPUE Year N 

Worm reefs, Indian River County, Florida 

(Data from Ehrhart et al., 2001) 38.86 2000 98 

Worm reefs, Indian River County, Florida 

(Data from Ehrhart et al., 1996) 6.28 1989 to 1995 190 

Nearshore reefs, Brevard County, Florida 3.52 2003 to 2004 20 

Nearshore reefs, Brevard County, Florida 1.25 2005 9 

Most adult female marine turtle species travel several hundred kilometers from foraging grounds to 

nesting beaches to lay eggs (Hirth, 1997). Nesting populations of turtles are believed to use nearshore 

waters predominantly as a corridor to the adjacent nesting beach, and green and loggerhead turtles have 

been observed mating close to shore (K. Holloway-Adkins, ECB, pers. observ.). However, adult 

loggerhead and green turtles have been frequently observed using OHB as rest areas during the nesting 

season in relatively large numbers, especially on reefs adjacent to Palm Beach County (Mortimer, 1995). 

There is no evidence to suggest that turtles laying subsequent clutches on east Florida beaches are 

foraging in adjacent waters during this period. 

High and low relief in NHB offers sheltered rest areas for marine turtles (Figure 6.5). Divers frequently 

observe loggerhead turtles resting under ledges and in shallow depressions on the reef 

(K. Holloway-Adkins, ECB, pers. observ.). Juvenile green turtles are easily captured, as they also rest 

under low relief areas and in shallow depressions in the reef (Makowski, 2004). Over 100 hawksbill 

turtles have been hand-captured from reefs in shallow (~2 m) and deep water (>18 m; Wood, 2006). 

Hawksbill turtles were more frequently captured on deeper reef areas, and all recaptured turtles were 

captured from deep reefs (L. Wood, Palm Beach Zoo, pers. comm.). 
Section 2 6-8 

Organismal Assemblages of East Florida Shallow Hardbottom 

Figure 6.5. Loggerhead (Caretta caretta) resting on offshore hardbottom, Breakers Reef, Palm Beach 

County, Florida. Photo by S. Hurley.

6.2.1 Green Turtles 

Juvenile green turtles are speculated to spend the first few years of their life in the pelagic (oceanic) zone, 

feeding on a variety of invertebrates and plants, and drifting with Sargassum racks along converging 

fronts (Carr and Meylan, 1980). However, efforts to confirm this hypothesis have been difficult 

(Witherington et al., 2006a). It appears that after approximately 2 to 5 years (at approximately 20 to 

25 cm), green turtles move into shallow coastal and estuarine waters (Hirth, 1997; Musick and Limpus, 

1997) (Figure 6.6). 

Figure 6.6. Juvenile green turtle (Chelonia mydas) over nearshore hardbottom in Palm Beach, Palm 

Beach County, Florida. Photo by D. Snyder.
Section 2 6-9 

Organismal Assemblages of East Florida Shallow Hardbottom 

At present, green turtles are the most frequently observed marine turtle species found associated with 

NHB from Brevard to Broward Counties. The presence of juvenile green turtles during surveys conducted 

during each season and observations of individual animals year-round suggest some animals 

continuously use NHB for several years. Where shallow NHB (intertidal and shallow subtidal waters) 

exists, juvenile green turtle densities were observed to be higher than they were in deeper, subtidal areas 

in Brevard (Holloway-Adkins, 2006), Indian River (Ehrhart et al., 2001), St. Lucie (Inwater Research 

Group, Inc., 2005), and Martin (Inwater Research Group, Inc., 2005) Counties. In areas where intertidal 

hardbottom was non-existent (Palm Beach and Broward Counties), juvenile green turtles utilized the first 

line of shallow reefs that paralleled the shore (Wershoven and Wershoven, 1992a; Makowski, 2004; 

Garrido, 2007). 

Ecological Associates, Inc. (2000) reported a substantial increase in the number of juvenile green turtles 

captured annually at the St. Lucie Power Plant after 1992, and their speculations as to why this has 

occurred include cooler water temperatures in northern regions, increased reef area providing a more 

abundant foraging resource, or an overall increase in the population of juvenile green turtles. None of 

these reasons have been substantiated to date; however, since their report (Ecological Associates, Inc., 

2000), the relative abundance of green turtles has still remained high on NHB. 

Mitochondrial DNA analysis conducted by Bagley (2003) and by Bass and Witzell (2000) indicate green 

turtles using NHB foraging grounds on the east coast of Florida showed high levels of genetic diversity. 

Bagley (2003) found 14 different haplotypes for green turtles captured over nearshore reefs in Indian 

River County. Haplotypes were from a mixed stock of nesting beaches from Florida, Mexico, Costa Rica, 

and Aves Island. 

Many researchers remark that the health condition of turtles captured or observed on NHB reefs typically 

is good and/or better than that of turtle populations residing in the Indian River and Mosquito Lagoon 

waters. Few animals from NHB study sites are afflicted with fleshy tumors (manifestations of 

fibropapillomatosis, or FP disease). None of the turtles captured in NHB waters in Brevard County have 

shown signs of the disease (Holloway-Adkins, 2005). Garrido (2007) did not sight any turtles on Palm 

Beach County reefs with the disease. 

Foraging sample analysis for turtles in Brevard, Indian River, Palm Beach, and Broward Counties indicate 

that a large percentage of the juvenile green turtle diet consists of red algae, with lower percentages of 

the diet from green and brown algae (Table 6.4). Relatively small amounts of invertebrate and inorganic 

food items (e.g., sand, shell, rock) were also found in foraging samples, and some samples contained 

pieces of plastic or other discarded marine debris. 

Juvenile green turtles have frequently been sighted in Palm Beach County, resting under ledges on 

shallow NHB (Jones et al., 2004). They were easily captured by Makowski (2004) while tucked under 

and around ledges west of Breakers Reef and on low relief areas and shallow depressions in the reef. 

6.2.2 Loggerhead Turtles 

Large juvenile loggerhead turtles recruit from oceanic environments into shallow coastal waters at >45 cm 

SCL (Witherington et al., 2006b). Loggerheads frequently have been captured during trawl events in 

areas north of the NHB region (i.e., Port Canaveral Inlet and shipping channel) (Schmid, 1995). They 

have also frequently been captured in the adjacent Indian River Lagoon system (Bresette et al., 2000). 

However, researchers infrequently sight or capture small size-class loggerheads in Brevard and Indian 

River County over shallow NHB habitats (Ehrhart et al., 2001; Holloway-Adkins and Provancha, 2005). 

Additionally, in St. Lucie County, juvenile loggerheads are systematically captured from the cool-water 

intake canals located in NHB (Ecological Associates, Inc., 2000). Juvenile loggerheads have been 

frequently observed foraging and resting on OHB in Palm Beach County (Mortimer, 1995). 

Loggerheads are considered to be omnivores. They forage on a wide variety of invertebrates, including 

salps, molluscs, and crustaceans. They tend to feed on more sessile organisms, many of which bury in 

soft bottom or sandy areas of reef and estuarine habitats. 
Section 2 6-10 

Organismal Assemblages of East Florida Shallow Hardbottom 

Table 6.4. Macroalgal species present in foraging samples from various population studies conducted on 

marine turtles on nearshore hardbottom habitat. ● denotes species presence. 

Macroalgal Forage Species Gilbert (2005) –

Indian River 

HollowayAdkins (2001)

– Indian River 

HollowayAdkins (2005) –

Brevard County

Jones et al. 

(2004) – Palm 

Beach 

Makowski 

(2004) – 

Palm Beach 

Wershoven and 

Wershoven 

(1992b) –

Broward 

Acanthophora muscoides ● 

Acanthophora spicifera ● ● ● 

Agardhiella subulata ● ● 

Amphiroa rigida ● 

Asteromenia (Fauchea peltata) ●

Bostrichia sp. ●

Botryocladia occidentalis ●

Botryocladia sp. ● 

Bryocladia cuspidata ● ● 

Bryothamnion seaforthii ● ● ●

Bryothamnion sp. ● ●

Caulerpa mexicana ● 

Caulerpa microphysa ● 

Caulerpa peltata ● 

Caulerpa prolifera ● ● ● 

Caulerpa racemosa ● ● 

Caulerpa taxifolia ● 

Centroceras clavulatum ● ● 

Ceramium sp. ● ● ● ● 

Chaetomorpha linum ● 

Chaetomorpha sp. ● ● ● 

Chondria dasyphylla ● 

Chondria sp. ● ● ● 

Chondrocanthus acicularis ● ● 

Cladophora catenata ● 

Cladophora sp. ● ● ● 

Codium sp. ● 

Dascycladus vermicularis ● 

Dasya pedicillata ● 

Dasya sp. ● 

Dictyopteris delicatula ● 

Dictyopteris sp. ● ● ●

Dictyospareia sp. ● 

Dictyota sp. ● ● ● 

Dudresyna crassa ● 

Erythrotrichia carnea ● 

Eucheuma nudum ● 

Gelidiaceae ●

Gelidiella sp. ●

Gelidium americanum ● ● ● 

Gelidium pusillum ● ● 

Gelidium sp. ● ● ●

Gracilaria armata ● 
Table 6.4. (Continued). 

Section 2 6-11 

Organismal Assemblages of East Florida Shallow Hardbottom 

Macroalgal Forage Species Gilbert (2005) –

Indian River 

HollowayAdkins (2001)

– Indian River 

HollowayAdkins (2005) –

Brevard County

Jones et al. 

(2004) – Palm 

Beach 

Makowski 

(2004) – 

Palm Beach 

Wershoven and 

Wershoven 

(1992b) –

Broward 

Gracilaria blodgettii ● 

Gracilaria mammillaris ● ● ● ● 

Gracilaria sp. ● ● ●

Gracilaria tikvahiae ● 

Gracilaria verrucosa ●

Halimeda discoidea ● 

Halimeda sp. ●

Halymenia floresia ● 

Hypnea cervicornis ●

Hypnea cornuta ●

Hypnea musciformis ●

Hypnea sp. ● ● ● ●

Hypnea spinella ● 

Hypnea valentiae ● 

Jania adherens ● ● 

Jania sp. ● ● 

Laurencia corallopsis ● 

Laurencia gemmifera ●

Laurencia poiteaui ● ● 

Laurencia sp. ● 

Liagora sp. ● 

Lomentaria baileyana ● 

Neomeris annulata ● 

Nitophyllum punctatum ●

Padina profunda ● ● 

Padina sp. ● ● 

Polysiphonia sp. ● 

Polysiphonia subtilissima ● 

Pterocladia sp. ●

Rhizoclonium sp. ● 

Rhodymenia sp. ● 

Sargassum sp. ● ● 

Schizothrix arenaria ● 

Schizothrix calcicola ●

Scinaia complanata ● ● 

Scinaia sp. ● 

Siphonocladus tropicus ● 

Solieria sp. ● ● 

Spyridia filamentosa ● 

Ulva sp. (Enteromorpha 

chaetomorphoides) ● 

Ulva sp. (Enteromorpha sp.) ● ● ● 

Ulva lactuca ● 

Ulva sp. ● ● ● 
Section 2 6-12 

Organismal Assemblages of East Florida Shallow Hardbottom 

6.2.3 Hawksbill Turtles 

Except for stranding data and sightings by SCUBA divers, little is known about the distribution of 

hawksbill turtles on NHB. Wood (2006) began studying E. imbricata on reefs off Palm Beach County in 

2004 and has captured over 100 hawksbills in water depths as shallow as 2 m and on deep reefs (>18 m) 

by SCUBA diving capture methods (L. Wood, in prep.). Based on SCL measurements (35.7 to 83.9 cm; 

mean 57.0 cm), this population is primarily composed of relatively large juvenile and subadult animals 

(Figure 6.7). Preliminary DNA analyses indicate a majority of these turtles are representative of Mexican 

stocks. Though not systematically studied until recently, SCUBA divers have reported an abundance of 

hawksbills in the area for decades (N. Rouse [deceased Divemaster, The Scuba Club, Inc., West Palm 

Beach, FL], personal dive logs; L. Wood, Palm Beach Zoo, pers. comm.).

Hawksbills are generally considered spongivores. However, a number of studies have shown their diet to 

be more varied than originally thought, and can include macroalgae, soft and hard corals, tunicates, 

jellyfishes and other small invertebrates that may or may not be associated with the sponges they 

commonly consume (Mayor et al., 1998; Leon, 2000; Leon and Bjorndal, 2002). Feeding observations 

conducted by L. Wood (Palm Beach Zoo, pers. comm.) reveal that hawksbills actively select and 

consume pieces and/or portions of sponges. This feeding behavior may play an important role in shaping 

the coral reef community and promoting the recolonization of sponges (Leon and Bjorndal, 2002). 

Jackson (1997) concluded that hawksbill turtles are keystone-like species in coral reefs systems. 

Hawksbill turtles are frequently approachable. They can be found out in the open, as well as under 

ledges or resting near reef structures (coral heads, barrel sponges, etc.). 

Figure 6.7. Juvenile hawksbill turtle (Eretmochelys imbricata) foraging on a tunicate wedged between the 

rocks on Nursery Reef, Pompano Beach, Florida (6 m water depth). A queen angelfish 

(Pomacanthus sp.) and gray angelfish (Holacanthus sp.) stood by, collecting scraps dropped 

by the hawksbill. Photo by K. Holloway-Adkins.
Section 2 6-13 

Organismal Assemblages of East Florida Shallow Hardbottom 

6.3 DISTURBANCE EVENTS AND TURTLES 

6.3.1 Shoreline Project Disturbances 

Juvenile green turtle abundance declined during surveys conducted between pre- and post-construction 

Broward County beach nourishment activities (Makowski and Kruempel, 2007). In one segment of the 

project area, green turtle abundance declined by 29.8% from pre-construction levels. In another 

segment, a 10.0% decline in turtle abundance was reported. Overall abundance level changes, however, 

were not statistically significant, and Prekel et al. (2007) suggest declines in abundance may be 

correlated with a temporary reduction in available macroalgae food resources. Third year 

post-nourishment biological monitoring surveys revealed that algae preferred by juvenile green turtles 

(based on foraging analysis from Wershoven and Wershoven, 1988 and Makowski et al., 2006a) were 

more abundant at 36-months post-construction. There appeared to be a temporary absence of turtles in 

the vicinity of a newly nourished section of beach in Indian River County in 2004. However, turtles 

returned to shallow subtidal areas shortly after water clarity was restored (K. Holloway-Adkins, ECB, pers. 

observ.). High turbidity levels from suspended sediments did not appear to affect foraging by juvenile 

green turtles, but excessive turbidity likely affects macroalgal food resources. 

6.3.2 Red Tide Disturbances 

Beginning in November 2007, red tide lingered onshore and offshore the east Florida coast for several 

weeks. FWRI (St. Petersburg) confirmed that red tide created from the dinoflagellate Karenia brevis, 

which originally bloomed off the west coast of Florida, was responsible for a major fish kill and several 

turtle deaths and sickness in Brevard and Volusia Counties. Over 30 juvenile green turtles were 

negatively affected by red tide events off the Brevard County coast (FWC personnel, pers. comm.). Many 

of the turtles that washed ashore during this event were transported to nearby rehabilitation facilities, and 

a few died. Loggerhead turtles succumbed to red tide toxicity and did not recover as well or as quickly as 

green turtles (M. Bauer, Volusia County Marine Science Center, pers. comm.). Many loggerhead turtles 

washed up dead or were transported live to facilities but did not recover. 

6.4 SUMMARY 

Data indicate that predominantly green and hawksbill turtles are the most frequently encountered species 

and the ones that assume relatively long-term residency on NHB. These data and observations suggest 

that the developmental life stages of juvenile green and hawksbill turtles, in particular, are intimately tied 

to the NHB habitat. The mean population size and size-class range suggest that the smaller mean size 

turtles (20 to 42 cm) are obligately tied to certain habitats within the NHB. At the very least, we are 

beginning to understand that juvenile green turtles are more susceptible to the debilitating 

fibropapillomatosis disease, which has been documented to occur in extremely high levels (50% or 

greater) in estuaries, bays, and sounds around the world (Herbst and Klein, 1995). Nevertheless, juvenile 

green turtles may not be obligately tied to NHB, but rather it might be to their advantage to have access to 

NHB for habitat. 

To date, only one area has been assigned critical habitat status for green and hawksbill turtles (Culebra, 

Puerto Rico). At present, marine turtle nesting beaches have been afforded more protection than 

developmental habitat for turtles. While beaches are important, the intermediate life stage (juvenile) 

might well be the most key component to the persistence and recovery of marine turtles (Bjorndal and 

Bolten, 2003). Until recently, we have attempted conservation of marine turtles with a focus on the 

individual species. A more effective approach may be to gain better insight into the functional roles of 

different life-history stages of marine turtles and adopt an approach that more strongly emphasizes 

ecosystem management with respect to all the habitats in which these animals are encountered 

(Costanza et al., 1997; Jackson et al., 2001; Bjorndal and Bolten, 2003). 
Section 3 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

Section 3 

Functional Attributes of Nearshore Hardbottom and 

Mitigation Alternatives 
Section 3 7-1 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

Chapter 7

Functional Ecological Attributes of Shallow Hardbottom

7.1 INTRODUCTION 

The overarching goal of this synthesis is to identify and provide supporting evidence of the important 

functional attributes of NHB, IHB, and shallow OHB. In this chapter, information in the preceding 

Chapters 3 through 6 on species and assemblage composition, interactions, and diversity will be 

examined in relation to the following priority issues: 

1. What are specific ecological functions of NHB? 

2. What can happen to the coastal ecosystem (broader than NHB) when NHB is lost due to 

nourishment? 

3. Is OHB habitat able to ecologically compensate for the loss of NHB? 

4. If the NHB habitat has specific properties and functions that cannot be compensated for by existing 

OHB habitat, what are the best mitigation alternatives for the loss of natural NHB? 

Richardson and Poloczanska (2008) examined traditional research approaches using functional 

groupings and individual species, preferring the latter. We recognize important roles for both approaches, 

depending on the starting objectives, and, given our objectives, will use both approaches according to 

topic. Advanced methods of examining functional roles using, for example, biological traits analysis 

(Bremner, 2008) may be of use in collective examination of NHB and deeper assemblages. 

The total number of reliably recorded species of invertebrates, fishes, algae, and other fauna from 

hardbottom from depths of 0 to approximately 4 m is at least 1,050 species along the mainland east 

Florida coast (Chapters 3, 4, 5, and 6). A total of 523 invertebrate species has been recorded to date. 

Among the most important invertebrates in terms of abundance and ecological function is the 

tube-building sabellariid polychaete worm Phragmatopoma lapidosa, which engineers structure utilized by 

at least 200 species of decapod crustaceans alone (Gore et al., 1978), many with direct or secondary 

trophic links to recreationally important fish species. However, the presence and significance of 

P. lapidosa declines south of Palm Beach County. The array of other significant invertebrates is large 

and includes Pione spp., Siderastrea spp., Menippe nodifrons, Pachygrapsus spp., Petrolisthes spp., 

Echinometra lucunter, and Synalpheus spp. The total number of fish species recorded is 257; several 

records await confirmation. The fish assemblage of NHB throughout the project region is a 

juvenile-dominated reef fish assemblage including grunts, damselfishes, wrasses, porgies, blennies, 

snappers, and other families of reef-associated fishes. 

The energetic functioning of NHB systems is in part driven by diverse macroalgal assemblages that 

include 275 species. The total number of algae and cyanobacteria recorded includes 277 identified 

species, with 341 total types and species, including unidentified species (Table 7.1). Of these, the 

rhodophytes (red algae) were most diverse, with 153 identified species. The most important species in 

terms of abundance and ecological function include Laurencia poiteaui, Caulerpa prolifera, Dictyota sp., 

Caulerpa sertularioides, and Caulerpa racemosa. 
Section 3 7-2 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

Table 7.1. Total numbers of family, genera, and species of macroalgae associated with nearshore 

hardbottom habitat from studies along the east Florida coast (see Appendix A, Table A.1). 

Group Division Family Genera Species 

Algae 

Chlorophyta 10 20 59 

Rhodophyta 30 62 153 

Phaeophyta 9 15 48 

Total algae 49 97 260 

Cyanobacteria 8 11 17 

Total number of species = 277; Total overall = 341 (includes unidentified types). 

7.2 ECOLOGICAL FUNCTIONAL GROUPING 

To structure the ecological complexity within shallow hardbottom, we organized the high diversity of 

species by identifying functional groupings that characterize the use of shelter and trophic resources and 

comparing patterns among primary functional groups, taxa, and other system components. Knowledge of 

the parts and processes that compose the functional attributes of shallow reef habitats in mainland east 

Florida is needed for management, but such efforts are frequently confounded by the morass of 

terminological holes involving the term function (Jax, 2005; Chapter 1, present study). 

Many studies have historically focused on two basic categories of functional attributes: 

1) structure-shelter and 2) feeding (trophic support). These groups can be useful as organizing steps in 

more detailed and measurable functional analyses such as traits analyses. Based on the literature on 

NHB and the literature on functions and functional groups, we identified at least 12 nominal functional 

group categories in Chapters 1 through 6 that apply to NHB, IHB, and shallow OHB assemblages: 

• Structure-shelter associated functional groupings. Examples include groups of species sharing one 

or more of the following attributes and associating with shallow hardbottom: 

− Habitat engineering functions (keystone framework builder); 

− Nesting and spawning site usage of NHB through OHB; 

− Structure as attachment resource for sessile organisms; 

− Settlement and post-settlement nursery areas; and 

− Juvenile and adult patterns of cross-shelf distributions.

• Trophic functional groupings. Food capture patterns, larger energetic fluxes and physical dynamics 

drive food web dynamics (e.g., inter-community nutrient exchange functions of carnivores and 

herbivores on NHB and OHB [Parrish, 1989]), such as the following: 

− Herbivory; 

− Carnivory; 

− Autotrophy; 

− Detritivory and omnivory; and 

− Cleaning symbioses. 

These 10 categories do not contradict the functional analysis framework associated with mitigation 

planning employed by the UMAM protocol (Continental Shelf Associates, Inc. et al., 2006). We also 

recognize that these are not entirely discrete groupings; there can be overlap among group processes 

and members, and also other attributes to emphasize. However, when synthetically evaluated for algae, 

invertebrates, fishes, and turtles, these groupings provide an heuristic structure that includes a majority of 

fundamental processes and functions of NHB, IHB, and shallow OHB. 
Section 3 7-3 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

7.2.1 Structure-Shelter Associated Functional Groupings 

Habitat and structure-associated processes influence most aspects of the life cycle of all shallow 

hardbottom organisms. The details of how spatial and structural habitat attributes influence plant and 

animal distributions have rarely been studied for shallow hardbottom systems of the region. Attributes of 

the nearshore seascape of mainland east Florida that are fundamental to nearshore marine habitats and 

their spatial relationships include the following: 

• NHB habitats are often separated from deeper hardbottom by several or many kilometers of relatively 

flat sand expanses; 

• North of Government Cut, Dade County, seagrass beds and mangroves are seen only in the lee of 

barrier islands and not in direct proximity to shallow hardbottom. These vegetated habitats occur 

inside the inlets of the study area and are typically not in direct proximity with shallow hardbottom as 

in the Florida Keys and Caribbean. Therefore, NHB is the only physical structure with raised relief 

(0.5 to 3.0 m emergent height) in shallow waters of mainland east Florida (sand bedforms may have 

relief of ≤0.5 m); 

• These nearshore areas are disturbance-mediated by depth and latitude. For example, higher energy 

nearshore wave environment exists north of the Bahamas wave shadow where more temperate 

coastal conditions also become more apparent and individual species adaptation to wider ranges of 

physical conditions is important; 

• Diversities of motile invertebrates correlate positively with the presence of P. lapidosa, while sessile 

organisms generally correlate negatively with the presence of P. lapidosa; and 

• The fish assemblages of shallow hardbottom in mainland east Florida are juvenile-dominated by 

species characteristic of coral reef fish assemblages. 

7.2.1.1 Habitat Engineers 

In terms of functional group considerations there is a prominent keystone component of hardbottom 

habitats along the Florida coast in the form of the habitat engineering polychaete P. lapidosa and the 

carbonate structures they rapidly build, commonly known as worm reefs or work rock. Worms of this 

sabellariid species select sediments from the water column and glue sand together to make sand tubes, 

forming substantial reefs in intertidal and shallow (<4 m) subtidal hardbottoms from Cape Canaveral to 

Key Biscayne in Florida. Their distribution continues southward to Santa Catarina, Brazil (Kirtley, 1994). 

Other representatives from their family (over 100 species) build extensive reefs that can be found in many 

locations throughout the world (reviewed by Kirtley, 1994; Pandolfi et al., 1998). 

In Florida, the structure provided by P. lapidosa worm reefs on top of coquina shellrock is a substantial 

contributor to biological diversity, supporting higher diversities and abundances of species, including 

hundreds of species of invertebrates (Gore et al., 1978; Nelson, 1989), than neighboring sand habitats 

and hardbottom without P. lapidosa colonization. Eight federally-listed and 15 state-listed species are 

associated with nearshore reefs off east Florida (USFWS, 1999). Additionally, green, hawksbill, and 

loggerhead turtles, primarily represented by juvenile and subadult stages, are recorded in association 

with shallow hardbottom habitats. In particular, worm reefs are important sources of food and shelter for 

juvenile green turtles (Wershoven and Wershoven, 1989; Ehrhart, 1992; Ehrhart et al., 1996; 

Holloway-Adkins, 2001; Makowski et al., 2006b). 

Worm reefs can also be important in the maintenance and persistence of beaches and barrier islands by 

retention of sediment, reduction of wave energy, grain sorting, and the progradation of beaches (Gram, 

1965; Kirtley, 1966, 1967; Multer and Milliman, 1967; Kirtley and Tanner, 1968; Mehta, 1973; Kirtley, 

1974; Pandolfi et al., 1998). P. lapidosa qualifies as a habitat engineer under criteria in Jones et al. 

(1994) and Coleman and Williams (2003) and has fast accretion rates (~0.5 cm per 24-hour period), with 

the potential to quickly create new structure with many microhabitat features, particularly subsequent to 

settlement events.
Section 3 7-4 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

7.2.1.2 Nesting and Spawning Sites 

Because many nearshore areas are expanses of sand in the absence of hardbottom, NHB may have 

substantial value as reference points for spawning activities of inshore fishes. Many fishes require 

three-dimensional structure as a reference point for coarse-scale aggregation and fine-scale behavior 

during spawning (Thresher, 1984). New information suggests that over 30 species may spawn directly on 

(nests) or near these reefs. Some are of substantial economic value; these include snook (Centropomus 

undecimalis), bigeye scad (Selar crumenophthalmus), and several herring species (Harengula spp.). 

Others, including small, cryptic species, especially blennies (Labrisomus nuchipinnis, Malacoctenus 

triangulatus, Parablennius marmoreus, and Scartella cristata) and damselfishes (Abudefduf saxatilis and 

Stegastes spp.) that deposit eggs directly on substratum, are common on NHB. 

Most species of invertebrates are broadcast spawners or they release gametes or larvae directly on or 

above shallow hardbottom. This particularly applies for the vast majority of sessile invertebrates in the 

project region, suggesting that at least 400 to 450 invertebrate species reproduce in association with 

shallow hardbottom. Reproduction in over 200 algal species also occurs in association with shallow 

hardbottom. 

7.2.1.3 Structure for Attachment 

The hard substrata associated with NHB is essential in providing a point of attachment for many sessile 

animals and algae. Without the substrate, organisms such as worm rock, sponges, crabs, many algal 

species, site-associated fish species, and others simply would not have the resources to exist in these 

subtidal areas. Along the east Florida coast, hardbottom facilitates worm rock recruitment, which in turn 

provides shelter for many motile invertebrates and fishes (Kirtley, 1966, 1974: Gore et al., 1978; USFWS, 

1999; McCarthy, 2001). Shelter is created not only by the external structure made by the worms but by 

the internal structure as well. The mounds created by P. lapidosa are relatively soft, and remarkable 

numbers of some invertebrates can be found within the worm rock, as some species of crabs, bivalves, 

and other worms are able to burrow in the rock to take shelter (Gore et al., 1978; Van Montfrans, 1981; 

Nelson and Demetriades, 1992). Other species such as snapping shrimp and sea cucumbers are seen 

using previously made crevices or holes. Overall, shelter inside the worm rock hosts flat worms, ribbon 

worms, sipunculans, many crustaceans, bivalves, and gastropods that would not exist in similar 

diversities or individual abundances in simple coquina rock hardbottom habitats. 

Hay and Sutherland (1988) emphasized the role of light in structuring rocky marine communities in the 

southeast U.S., with well-lit structure often dominated by algae and unlit areas by invertebrates. This 

pattern may generically apply to shallow hardbottom of mainland east Florida, but it is not yet tested. The 

high diversities of algae would not exist without substrate for over 200 species to attach to. A variety of 

opportunities for intertidal and subtidal research into rocky intertidal processes involving invertebrates, 

algae, and their interactions remain relatively unexamined in association with shallow hardbottom of east 

Florida. 

In addition to providing essential functions for numerous fishery species, bioerosion of hardbottom 

provides a source of new microhabitat and sand on the continental shelf (Riggs et al., 1985). Boring and 

burrowing shrimp and bivalves chemically or mechanically excavate holes, eventually weakening the 

rock; clionid sponges and rock-boring urchins do this as well. This habitat engineering manifests itself in 

differing ways. For example, crabs bore into worm mounds, and the synergistic effect of these boring 

organisms is high when coupled with natural disturbances such as wave activity. These all contribute to 

worm reef degradation and an increase in structural complexity by late summer/early fall. This process 

also enhances the structural complexity of hardbottom outcrops, promoting diversity of reef microhabitat 

structure. 
Section 3 7-5 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

7.2.1.4 Settlement and Post-settlement Nursery Areas 

Invertebrates 

It is probable that most of the motile invertebrates that settle on intertidal and shallow hardbottom along 

the coast ultimately die in the same areas that they settle. Many species that thrive in these shallow 

habitats (P. lapidosa, Pione lampa, and Siderastrea radians) become sexually mature very young and 

reproduce continuously (Soong, 1993; McCarthy, 2001; Schönberg, 2002; McCarthy et al., 2003). Most 

research on gastropods and crustaceans suggests that animals settle there and live a maximum of 2 to 

3 years because of the ephemeral nature of these habitats (Gore et al., 1978; McCarthy, 2001; 

Watanabe, 2002; Watanabe and Young, 2006). For example, the oyster drill Stramonita (Thais) 

haemostoma has a population size distribution that gets progressively larger from fall to winter (but never 

as big as their oyster drill counterparts within the Indian River Lagoon) until seasonal disturbance results 

in high mortality by late summer (Watanabe, 2002; Watanabe and Young, 2006). However, they have 

adapted by becoming reproductively active faster than their lagoonal counterparts so that they can lay 

eggs before they die and produce a new cohort seemingly to maintain the population. Consequently, 

these new recruits settle in high numbers and shift the population distribution size back to smaller modal 

sizes. 

High mortality of P. lapidosa mounds is common in some NHB areas of east-central Florida and increases 

during the summer as a result of the effects of several disturbance agents (McCarthy, 2001). In the early 

summer, some individuals at the tops of intertidal mounds perish, leaving the tops susceptible to decay 

and colonization by other sessile organisms. It is likely that this mortality is caused by desiccation and/or 

heat stress from extreme summer temperatures. By the late summer and early autumn, wave activity 

from hurricanes results in maximum physical disturbance to sabellariid reefs. A large percentage of both 

intertidal and subtidal reefs are severely damaged at this time. Intertidal worms are more susceptible to 

physical destruction of their colonies, whereas subtidal worms get smothered by sand but the sand reef 

remains intact. In addition, post-hurricane deposition of muddy sediment can kill attached invertebrates 

and prevent settlement (CSA International, Inc., 2007; Miller and Kosmynin, 2008). 

Almost simultaneously with peaks in lethal disturbance, however, larvae of P. lapidosa arrive in large 

numbers to renew the reef by massive recruitment in cracks, atop, or on edges of adult mounds 

(McCarthy, 2001). Generally, recruitment occurs at the highest levels in the intertidal versus subtidal 

zone (Figure 7.1). McCarthy (2001) suggests that subtidal recruitment may be enhanced during periods 

of increased wave activity. Preliminary results from an ongoing study suggest that among five intertidal 

locations along a 218-km stretch of east Florida (Satellite Beach, Stuart, Coral Cove, Palm Beach, 

Boynton Beach), during the years from 2006 to present, P. lapidosa recruited at the highest levels in 

Stuart, followed by Satellite Beach. However, at all five locations, peak larval recruitment appears to 

occur during the fall-winter time periods. Consequently, recruitment during this time results in new 

low-lying reefs that are highly resilient and will eventually restore the structure of the reefs. As 

disturbance lowers adult abundance and creates new settlement space, new individuals potentially can 

arrive in sufficient numbers to restore the populations. Therefore, local populations may remain at fairly 

high abundances year after year while experiencing moderately high mortality from various agents of 

disturbance (McCarthy, 2001; Figure 7.1), a pattern seen in some intertidal organisms in the Pacific U.S. 

When this P. lapidosa colonization information is integrated with those of other researchers (Gilmore, 

1977; Gilmore et al., 1981; Lindeman and Snyder, 1999), links are suggested between the seasonal cycle 

of sabellariid reef expansion and degradation, and the occupation of those reefs by juveniles and adults 

of other organisms. 
Section 3 7-6 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

Recruits m-2 d-1

1e-1

1e+0

1e+1

1e+2

1e+3

1e+4

1e+5

Month and Year

J J ASOND J FMAMJ J ASONDJ FMAMJ J ASONDJ FMAM

1

10

100

1000

10000

Intertidal habitat

Subtidal habitat

1997 1998 1999 2000

Figure 7.1. Seasonal larval recruitment of the sabellariid worm Phragmatopoma lapidosa in intertidal and 

subtidal nearshore hardbottom habitats off Boynton Beach, Palm Beach County, Florida. 

Mean values (± standard deviation) of recruitment are plotted on log scales. All subtidal sites 

covered with sand after May 1999 (From: McCarthy, 2001). 
Section 3 7-7 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

Some biological information on ephemeral exposure of NHB is available in regards to short-term 

P. lapidosa tolerance to sand covering. These worms typically have some tolerance to limited sand 

covering for short periods of time, up to 72 hours in various studies (Main and Nelson, 1988; Sloan and 

Irlandi, 2007). After this time period, most individual worms die. Mortality occurs earlier during warmer 

periods. Scleractinian corals (e.g., Siderastrea radians) have been recorded growing on seasonally 

buried NHB (V. Kosmynin, FDEP Bureau of Beaches and Coastal Systems, pers. comm.). Care should 

be exercised in averaging NHB exposure over multiple years for beach nourishment projects. The 

majority of beaches and nearshore sedimentary systems have been exposed to several renourishments 

or have been indirectly affected by long-shore drift. To best capture natural situations and away-fromnourishment effects, photoimagery is best taken right before dredge and fill events when aerial resources 

are available and visibility allows (V. Kosmynin, FDEP Bureau of Beaches and Coastal Systems, pers. 

comm.). 

On hardbottom deeper than 4 m, the degree of disturbance is likely less; there is evidence that this may 

correlate with the presence of longer-lived invertebrate species (D. McCarthy, Jacksonville University, 

Jacksonville, FL, pers. observ.). There appears to be a higher number of sessile species that take longer 

to become sexually mature and reproduce seasonally. It is unclear how much movement there is of 

motile invertebrates among hardbottoms within this depth range. However, spiny lobster clearly move 

long distances among hardbottom habitats, as seen in the Florida Keys and Bahamas (Kanciruk and 

Herrnkind, 1978) (see Section 7.2.1.5). 

Fishes 

In terms of ecological function, several lines of evidence suggest that NHB reefs may serve as significant 

settlement areas in shallow marine waters from Broward through Brevard Counties. The following 

summary of fishes is based on information available for mainland southeast and east central Florida 

(Gilmore et al., 1981; Lindeman, 1997a; Lindeman and Snyder, 1999; Baron et al., 2004), with some 

papers including life stage-specific abundance data. Pooled early life stages can represent over 80% of 

the total individuals at sites censused (Figure 5.6). Eight of the top 10 most abundant species were 

consistently represented by early stages. Use of hardbottom habitats was recorded for newly settled 

stages of more than 20 species. Some species have characteristic settlement microhabitats in which 

they can differentially aggregate while undergoing transformation from epibenthic larvae into demersal 

juvenile stages. Microhabitat adaptation in newly settled drums (Pareques spp., Sciaenidae) can involve 

the occurrence of triangularity, for example in the frontal and lateral profiles of individual larvae, the 

crevices between rock and coarse sand they typically utilize, and the outer boundary shapes of schools 

that settlers can form (Figure 7.2). 

Figure 7.2. Newly settled drums (Pareques species) commonly associate with ledges and triangular 

cavities at the base of shallow hardbottom structures, such as in this photograph taken from 

2 m depth, Singer Island, Palm Beach County, Florida. Photo by D. Snyder.
Section 3 7-8 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

The presence of more juvenile stages than adults does not guarantee a habitat is a valuable nursery 

(Adams et al., 2006). Rapid decays in the benthic or planktonic survival of early stages of marine fishes 

are common demographic patterns (Shulman and Ogden, 1987; Jones, 1991), ensuring that if 

distributions are homogeneous, all habitats will have more early stages than adults. Settlement and early 

life stage utilization of shallow hardbottom has been studied in more detail in recent years, in part due to 

artificial reef placement in IHB and OHB depths. Despite substantial sampling efforts in deeper waters, 

many species of grunts are not encountered as early stages in midshelf or deepter reef habitats 

(e.g., Jordan et al., 2004). Adults are often infrequent or absent from the shallow habitats. There is 

considerable evidence for habitat segregation among life stages of some species, with the earliest stages 

of many species of grunts and snappers often using the shallower habitats (Starck, 1970; Dennis, 1992; 

Lindeman et al., 1998; Jordan et al., 2004). Similar ontogenetic differences in both distribution and 

abundance exist for diverse species from other families that utilize NHB habitats. Based on available 

information, we estimate that at least 40 fish species utilize NHB as a primary or secondary larval 

settlement area, many (but not all) with seasonal peaks in warmer months (Lindeman and Snyder, 1999; 

Baron et al., 2004). 

In mainland east Florida, NHB may also serve as intermediate nursery habitat for late juveniles emigrating 

out of estuaries (Vare, 1991; Lindeman and Snyder, 1999). This appears to also apply to some fishes 

from hardbottom to the north. Nearshore and inner shelf hardbottom areas north of Cape Canaveral can 

also serve as important settlement and nursery habitat for immigrating larvae of coastal fish species. 

Powell and Robbins (1998) collected larvae from 22 reef-associated families adjacent to hardbottom 

habitat in Onslow Bay, North Carolina. Tomtate (Haemulon aurolineatum), white grunt (H. plumieri), 

snappers (including vermilion snapper, Rhomboplites aurorubens), black sea bass (Centropristis striata), 

bank sea bass (C. ocyurus), sand perch (Diplectrum formosum), spottail pinfish (Diplodus holbrooki), and 

whitebone porgy (Calamus leucosteus) were commonly collected (SAFMC, in prep.). These species may 

have been spawned in Onslow Bay in somewhat deeper water and recruited locally to nearshore 

hardbottom (Powell and Robbins, 1998; SAFMC, in prep.). 

7.2.1.5 Juvenile and Adult Cross-shelf Distributions 

The vast majority of invertebrate species do not migrate or shift habitats from shallow hardbottom. Some 

motile exceptions may include portunid crabs, spiny lobster, and squids, but there is not a great deal of 

invertebrate migration between NHB and neighboring habitats. The polychaete P. lapidosa generally has 

higher abundance intertidally than subtidally along east Florida hardbottoms (Figure 7.1). It is 

moderately abundant in subtidal NHB and somewhat common in IHB. Generally, P. lapidosa is in very 

low abundance or not encountered in nearshore OHB along the east Florida coast. 

Spiny lobster, primarily known from NHB in Indian River and St. Lucie Counties, clearly move long 

distances from shallow to deep habitats, as seen in the Florida Keys and Bahamas (Kanciruk and 

Herrnkind, 1978). Spiny lobsters have been observed moving along 4 m reef at Boynton in chains of 

~15 (D. McCarthy, Jacksonville University, Jacksonville, FL, pers. observ.). Within that area of the coast, 

there were probably hundreds on the move heading northward. Mass migrations (referred to as “crawls” 

by locals) by large numbers of spiny lobsters occurs typically in late summer early fall along the coast of 

southeast Florida. Single-file lines of individual lobsters move presumably from offshore areas toward the 

shore where they seek cover around any structured object (natural or artificial). Kanciruk and Herrnkind 

(1978) determined that mass migration in the Bahamas was triggered by the onset of cold fronts. At least 

one mass migration in 1980 off Palm Beach, Florida was associated with a cold water upwelling event 

(D. Snyder, CSA International, Inc., pers. observ.). 

Damselfishes and cryptic hardbottom fish species do not engage in ontogenetic habitat shifts. Juveniles 

of schooling species (surgeonfishes, parrotfishes, chubs) may migrate away from NHB to deeper 

structural habitats with maturation, but depth-specific information is largely unavailable among these 

habitats. The ability to successfully make such migrations will correlate in complex manners with the 

degree to which cross-shelf habitat corridors are available and, also, drivers associated with potential 

ontogenetic shifts in food needs. Cryptic resident species do not appear to migrate further offshore with 

development, displaying no cross-shelf ontogenetic migrations. They are life-long residents of the NHB, 
Section 3 7-9 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

IHB, or shallow OHB structure that they settle on and display feeding modes from herbivory to probable 

piscivory. A variety of other species (e.g., wrasses, grunts, snappers, sciaenids, mullids, S. rubripinne) 

may sample other habitats with age and eventually emigrate; presumably survival will be very dependent 

on the habitat and predator seascape, both while traveling and eventually on entering new habitat. 

Juvenile marine turtles off Florida coastal waters are predominantly found associated with NHB habitats. 

Recapture data and tracking information from population studies suggest that turtles spend several years 

in neritic habitats (Ehrhart et al., 2001; Makowski et al., 2006b). Juvenile green turtles have been 

recaptured on the same NHB reef locations more than 9 years later (Ehrhart, unpubl. data). Studies in 

NHB habitat indicate that juvenile green turtles grow slower in NHB versus estuarine habitats (Kubis, 

2003); however, the costs of slow growth may be offset by relatively lower predation levels due to higher 

structural complexity offering better protection from predators. 

7.2.2 Trophic Functional Groupings 

7.2.2.1 Herbivory 

Invertebrate herbivory on macro- and micro-algae may be high among the diverse array of plants 

associated with shallow hardbottom. In various temperate rocky intertidal studies, over 50 species of 

shallow hardbottom or rocky intertidal invertebrates have been reported to feed on macro- and 

microalgae (Appendix D, Table D.1). Some of the same macroalgal genera are found on Florida NHB 

(Acanthophora, Bryopsis, Ceramium, Chondria, Codium, Dictyota, and Sargassum) and are likely grazed 

on by fish and invertebrate herbivores. Vertebrate grazing analyses from juvenile green turtle foraging 

samples (Holloway-Adkins, 2001) and herbivorous fishes (Randall, 1967) indicate Ceramium sp., Ulva

sp., Polysiphonia sp., Lyngbya sp., Centroceras sp., and Dictyota sp., are primary food items for these 

animals (Appendix D, Tables D.2 and D.3). 

Along east Florida hardbottom there are a number of echinoids that may be important herbivores. The 

most significant are Echinometra lucunter, Tripneustes ventricosus, and Arbacia punctulata. They are 

generally most abundant to the north of Palm Beach County at depths in excess of 1 m and occur in 

highly variable densities that can range as high as ~25 individuals /m2

 and likely exert substantial 

herbivory affects on marcoalgae and sessile invertebrates in the areas where they occur. Other 

echinoids such as Diadema antillarum, Lytechinus variegatus, and Eucidaris tribuloides are herbivores 

but probably do not exert the same influence over macroalgal communities because of their generally low 

abundance. 

Many reef fish herbivores occur in association with NHB and include the surgeonfishes (Acanthuridae), 

parrotfishes (Scaridae), porgies (Sparidae), and chubs (Kyphosidae); most of these families are 

represented largely by juvenile life stages on NHB, but can occur as adults (Lindeman and Snyder, 1999; 

Baron et al., 2004). Non-schooling herbivores on NHB include the damselfishes (five species), filefishes 

(Monacanthidae), spiny puffers (Diodontidae), presumably all gobiids, and several blennids (Randall, 

1967). In NHB environments, herbivores represented relatively few fish species (Figure 5.11), but these 

can be locally abundant and may have some impacts on macroalgal biomass, though this is unstudied. 

There can be substantial ontogenetic variation in feeding among some species. For example, the silver 

porgy (Diplodus argenteus), a very abundant sparid on NHB, is probably omnivorous like its closely 

related congener, the spottail pinfish (D. holbrooki). The latter species was found to eat both algae and 

macroinvertebrates by Hay and Sutherland (1988) in a study of rocky jetty-type structures in Murell’s Inlet, 

South Carolina. Carr and Adams (1972) documented that D. holbrooki fed upon ectoparasites and were 

therefore considered to be cleaners. Juveniles of D. argenteus have been observed cleaning larger 

fishes in NHB environments (D. Snyder, CSA International, Inc., pers. observ.). 

Juvenile green turtles preferentially feed on several algal species from a larger pool of available algae 

(Appendix D, Table D.3). High wave energy not only strongly influences the production of macroalgae, 

but also can reduce grazer accessibility, especially in intertidal and shallow subtidal sections of NHB and 

IHB. This species may be more abundant where competition from herbivorous fish and invertebrate 
Section 3 7-10 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

species is reduced (e.g., the northern project region with cooler waters). Macroalgal resources, in turn, 

may be more abundant for several reasons, including cooler water temperatures, higher nutrient levels, 

and, potentially, a lower abundance and diversity of grazers. 

7.2.2.2 Carnivory 

Invertivory 

Crustaceans are often very important predators upon invertebrates within shallow hardbottom food webs 

along the east Florida coast. Some crustaceans can also feed on small fishes. Gore et al. (1978) 

postulated a possible food web for sabellariid colonies on the east Florida coast where the worms form 

the basis of the food web on which many species of reef-dwelling crabs forage (Figure 7.3). 

Figure 7.3. Food web among six common decapod crustaceans from worm reefs. Numbers on lines 

indicate the percent of observed prey material in the guts of all individuals examined per 

species. Dashed lines indicate production from worm reef (Phragmatopoma lapidosa) 

(From: Gore et al., 1978). 

There is a high diversity and abundance of infaunal invertebrates within soft sediments adjacent to east 

Florida NHB. Within the intertidal and shallow subtidal zones, there are various crustaceans, molluscs, 

bivalves, and polychaetes that can be very abundant. Mole crabs (Emerita talpoida) and various Donax 

spp. are often most conspicuous. However, several species of polychaetes such as Nephtys bucera and 

Scolelepis squamata are common in these habitats as well. Progressing into deeper sand bottom areas 

(~3 m water depths), the polychaetes Dispio uncinata and Paraonis sp. and the bivalve Tellina sp. are 

very dominant. However, amphipods, caridean shrimp, portunid crabs, gastropods, and bivalves can also 

be encountered. Some threatened coastal bird species feed on intertidal invertebrates and can exchange 

energy between the upper beachface and the NHB, or the coastal ocean and NHB. Species include 

herons, egrets, plovers, brown pelicans, and others (Bishop et al., 2006). 

Many demersal fish species may migrate further into adjacent sand areas in search of food as they 

mature, and they can exchange large amounts of nutrients in these often nocturnal migrations (Parrish, 

1989). Meyer et al. (1983) provides a specific example of intercommunity energy exchange and growth 
Section 3 7-11 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

impacts for grunts in the U.S. Virgin Islands. Older life stages of grunts and other species on NHB may 

be engaging in sand feeding migrations at night, exchanging energy between reef and softbottom 

environments. In addition, coastal pelagics can transport energy into and out of the system via transient 

usage of NHB and IHB structure. 

The majority of the approximately 200 fish species recorded from NHB through OHB in east Florida feed 

on invertebrates, first as zooplanktivorous larvae and via an explosion of diversity in invertebrate 

consumption with maturation among reef fishes (Figure 7.4). There are often large variations in prey 

composition among differing life stages. Detailed studies on trophic ecological process of shallow 

invertebrates and fishes of Florida NHB 

are lacking, therefore constraining 

analyses. 

Planktivory and Suspension Feeding 

A number of sessile invertebrates 

encountered along east Florida 

hardbottoms are suspension feeders. 

Within the intertidal sections of the NHB, 

P. lapidosa and several barnacle species 

are the most commonly encountered. 

From subtidal NHB to OHB, P. lapidosa

can be abundant in shallower depths, but 

a number of sponge and tunicate species 

become more prevalent. Additional 

suspension feeders encountered in these 

habitats are sabellid and serpulid 

polychaetes, several species of bivalves, 

and decapods such as porcellanid crabs. 

At least 30 fish species associated with NHB can be classifed as planktivorous. Common planktivores 

found on NHB including herrings (Harengula spp., Opisthonema oglinum), sardines (Sardinella aurita), 

and scads (Decapterus spp., Selar crumenophthalmus) tend to form large schools and frequently move 

within local areas. Many fishes are planktivorous as newly settled or early juvenile individuals 

(haemulids, reef sciaenids) but change diets with ontogeny. Silver porgy and spottail pinfish are 

examples of species that shift from planktivory as young to omnivory as adults (Carr and Adams, 1972). 

Grunts such as French grunt and tomtate are obligate planktivores as newly settled individuals but 

gradually switch to demersal carnivory as they grow. Observations on NHB in Palm Beach County 

indicate that larger juveniles of the these species will opportunistically feed on plankton when directional 

currents are strong and presumably carrying greater quantities of plankton over the NHB. Given the high 

proportion of early stage fishes found on NHB, planktivory represents an important trophic element of the 

local system that is subsidized by the import of plankton from the broader scale coastal waters. At 

present nothing is known regarding the diversity, composition, or temporal occurrence of the planktonic 

prey base in NHB environments. 

7.2.2.3 Autotrophy 

Cyanobacteria and macroalgae, the primary producers in marine ecosystems, convert sunlight and 

nutrients into assimilable energy used by primary consumers. These photosynthetic cohorts specialize in 

the ability to convert abiotic material (carbon) and light into food. Macroalgae utilize both nutrients and 

carbon dioxide and are able to fix nitrogen, and many species of cyanobacteria also can fix nitrogen 

(Vermaas, 2001). Primary animal autotrophs in this system are the zooxanthellae-bearing 

sceleractinians. In addition to autotrophic production, these species have multiple ecological functions as 

shelter and direct food resources (Banks et al., 2008). While primary production is the foundation of the 

trophic food pyramid, the importance of maintaining balanced trophic levels in marine systems, especially 

coral reefs, is high (e.g., Paddack et al., 2006). Overgrowth of coral reefs by macroalgae due to 

Figure 7.4. Gray triggerfish, Balistes capriscus, pursuing 

invertebrate prey, Breakers Reef, 

approximately 6 m depth, Palm Beach, 

Florida. Photo by D. Snyder.
Section 3 7-12 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

increased nutrient input or a significant reduction of primary consumers by overfishing can have large, 

well-documented impacts on coastal food webs (Jackson et al., 2001). 

There may be a generally higher biomass of macroalgae in the more northern sections of hardbottom 

along the east Florida coast. The exact reasons for this trend are unclear. The Florida Current may 

contribute to this trend because its coastal influence decreases towards the north. Consequently, there 

are cooler waters and possibly higher upwelled nutrients in the more northern hardbottom areas along the 

coast. More nutrients may provide important resources to increase algal biomass. Additionally, towards 

the south there may be a higher diversity and abundance of herbivorous fishes that may affect macroalgal 

production in those areas, thus creating space for otherwise substrate-limited organisms (i.e., corals). 

These and other alternatives are ripe for empirical investigation. 

7.2.2.4 Detritivory and Omnivory 

Many detritivores in association with NHB are omnivorous. In marine food webs, detritivores play 

important roles as recyclers in ecosystem energy flow and biogeochemical cycles. A number of motile 

invertebrates found on shallow hardbottom are known to feed on detritus, although no known studies 

have specifically investigated invertebrate detritivory on NHB. Most known invertebrate detritivores found 

on Florida NHB are polychaetes or crustaceans, although some gastropods and echinoderms are likely to 

feed on detritus as well. The Gore et al. (1978) trophic web for worm reef areas suggested that the highly 

abundant crustacean Pachycheles monilifer was abundant in shallow hardbottom habitats because of its 

omnivorous diet that includes detritus in suspension. They also suggested that various amphipods, 

isopods, molluscs, and sponges gain nutrition from suspended detritus. 

There are a number of motile invertebrates and some fish species on NHB that can be considered 

omnivores. Most omnivorous invertebrates are crustaceans, and several are very abundant. A number 

of xanthid, portunid, and hermit crabs and shrimp can be omnivorous. Slipper, spiny, and spotted spiny 

lobster are omnivores that are also common in subtidal NHB, IHB, and OHB. Some molluscs (e.g., a few 

gastropods and cephalopods) can be omnivorous as well. As adults, omnivorous species of fishes 

include chubs (Kyphosus species), filefishes and puffers, all gobiids, and several blennioids. When whole 

ontogeny is considered, some species are relatively omnivorous, feeding on plankton not only as larvae 

but sometimes for a week or more after settlement (e.g., haemulids, some sciaenids, pempherids). 

7.2.2.5 Cleaning Symbioses 

Cleaning behavior is defined as the removal by a cleaning organism of ectoparasites or injured tissue 

from a host or client organism. In marine environments, shrimp and fishes are the primary taxa that 

engage in cleaning behavior. Despite the many examples of cleaning behavior, only a few species are 

obligate cleaners, whereas most of the facultative cleaners do 

so only as juveniles (e.g., Hobson, 1971; Poulin and Grutter, 

1996; Sazima et al., 1999). Hobson believed that only certain 

individuals of some species clean and that cleaning behavior is 

widespread among small species that pick organisms from the 

substrate or water column. 

Examples of fish species recorded from shallow hardbottom 

that act as facultative cleaners include the young of leatherjack 

(Oligoplites saurus) (Lucas and Benkert, 1983), spottail pinfish 

(Diplodus holbrooki) (Carr and Adams, 1972), porkfish 

(Anisotremus virginicus) (Longley and Hildebrand, 1941), 

angelfishes (Pomacanthus spp. and Holacanthus spp.) 

(Brockmann and Hailman, 1976), and the wrasses (Feitoza et 

al., 2002). Cleaning behavior can frequently be observed on 

NHB, with cleaners including juvenile angelfishes, wrasses, 

gobies, porkfish, and silver porgy (D. Snyder, CSA 

International, Inc., pers. observ.). Figure 7.5 shows a juvenile 

Figure 7.5. A juvenile French angelfish 

(Pomacanthus paru) 

cleaning an adult gray 

triggerfish (Balistes 

capriscus) in 4 m depth off 

Palm Beach, Palm Beach 

County, Florida. Photo by 

D. Snyder.
Section 3 7-13 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

angelfish cleaning a gray triggerfish on NHB near Palm Beach, Florida. Baron et al. (2004) included nine 

species of known cleaners during their Broward County study. Lindeman and Snyder (1999) encountered 

six known cleaning species in the Jupiter area. Shrimp known to be cleaners, including Stenopus 

hispidus, S. scutelous, Lysmata spp., and Periclimenes pedersoni, do inhabit NHB (D. Snyder [CSA 

International, Inc.] and D. McCarthy [Jacksonville University, Jacksonville, FL], pers. comm.); however, 

none were recorded during the sampling programs discussed in Chapter 4. 

7.3 LATITUDINAL COMPARISONS 

Sessile invertebrate communities change noticeably with latitude along the east Florida coast. In 

intertidal hardbottoms to the north, dominant invertebrates are bryozoans, sabellariid polychaetes, sea 

anemones, and tunicates (Chapter 4). North of Palm Beach County, there is an increase in abundance 

of P. lapidosa, both intertidally and subtidally. Throughout tidal pools within intertidal areas of these 

latitudes, the starlet coral Siderastrea radians, two species of zoanthids, and several species of solitary 

anemones can be encountered. Within the lower intertidal and shallow subtidal zones, the most 

conspicuous sponge is the rock-boring sponge Pione lampa. In the more northern counties that have 

subtidal hardbottom (i.e., Brevard and Indian River), sponges, two species of gorgonians (Leptogorgia 

virgulata and L. hebes), and one species of telestacean (Carijoa [Telesto] riisei) can be encountered. In 

this area, the most significant hard corals are Siderastrea and the Oculina species. However, there are 

occasional occurrences of Cladocora arbuscula, and grouped polyps of Phyllangia and Astrangia spp. 

(Brooke and Young, 2005). Within Martin County, the St. Lucie Inlet Reef appears to be the northernmost 

extent of several of the reef-building corals that are known biodiversity contributors. 

To the south, the diversity and abundance of anthozoans increase on subtidal hardbottoms (SFCREMP, 

2007; Beal, unpublished data). The geological origins of Broward’s NHB have some different 

characteristics than northern areas (Moyer et al., 2003; Banks et al., 2008). Unlike counties to the north, 

Broward County possesses a nearshore ridge complex inside the inner reef line that has a complicated 

array of depth strata. The shallowest prominent scleractinians in the 3 to 7 m depth range include 

Solenastrea bournoni, Diploria clivosa, Dichocoenia stokesii, and the staghorn coral, Acropora 

cervicornis (Moyer et al., 2003; Vargas-Angel et al., 2003; SFCREMP, 2007). The potential biological 

significance of these latitudinal differences for shallow hardbottom communities has not been formally 

evaluated for any invertebrate species to date. 

Differences among ichthyofaunal assemblages are suggested between the southern and northern areas 

of mainland southeast and east-central Florida in terms of the most abundant species (Chapter 5), but 

information remains relatively limited. Causes of the differences can include 1) differing amounts and 

types of shallow hardbottom among sections of southeast Florida and east-central Florida and 

2) differences in northern geographic distributional boundaries among some highly abundant species. 

There are notable differences in abundance between three of the most abundant species in the Jupiter 

area in comparison to the Fort Lauderdale area. Only one black margate out of >72,000 individuals was 

recorded in Fort Lauderdale (Baron et al., 2004), while 636 black margates out of 10,491 individuals were 

censused in Lindeman and Snyder (1999). Also, 43 sailors choice, Haemulon parra, were recorded in 

Baron et al. (2004), while 1,326 H. parra individuals were recorded in Lindeman and Snyder (1999). The 

hairy blenny was the most abundant cryptic hardbottom species in the Jupiter surveys, 8% of the overall 

individual total (Lindeman and Snyder, 1999); none were recorded in transect and point-counts in Fort 

Lauderdale (Baron et al., 2004). Numbers of the abundant porgies, Diplodus spp., were also much lower. 

Seasonality of sampling will be important as comparative information is examined further. 

The latitudinal distribution of marine turtles associated with NHB is unclear at this time. General patterns, 

however, indicate that loggerhead turtles are sparsely present in northern NHB (Brevard and Indian River 

Counties) and moderately present on deeper reefs in southern NHB (Palm Beach County). Visual 

transect survey data indicate that juvenile green turtles are more abundant in intertidal and shallow 

subtidal areas than in deeper subtidal areas, and hawksbill turtles are infrequently encountered north of 

Palm Beach County. 
Section 3 7-14 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

7.4 DISTURBANCE ECOLOGY 

Disturbance has been defined in several ways (see Sousa, 2001 for a review). It is often defined as 

damage or mortality that is caused by some external agent or force. This agent can be either physical 

(e.g., wave activity) or biological (e.g., predation) (Sousa, 2001). Disturbance may be lethal, resulting in 

the death of the animal, or it may a sublethal stress that reduces the potential for growth or reproduction 

of an organism. Obviously, any disturbance can affect reproduction if it kills the animal before it has the 

chance to reproduce. Sublethal disturbance can also influence reproduction by affecting the available 

energy an animal has for reproduction. Several correlative studies on marine invertebrates have shown 

negative relationships between fecundity and external stress factors such as sedimentation, wave activity, 

and turbidity (Kojis and Quinn, 1984; Jokiel, 1985; Rinkevich and Loya, 1987; Tomascik and Sander, 

1987; Szmant and Gassman, 1990; Van Veghel and Bak, 1993; Ward, 1995). 

Nonlethal disturbance might affect gamete production by diverting energy to regeneration of body parts 

(Lawrence and Vasquez, 1996). Such disturbance is known to occur to individuals by a variety of 

mechanisms, including unsuccessful predation attempts, trampling by humans or other animals, or the 

impact of objects propelled by waves (see Sousa, 2001 for a review). Stress caused by changes in 

several environmental factors has been implicated in numerous studies of spawning and/or gametogenic 

synchrony in marine invertebrates (Young, 1945; see Giese and Kanatani, 1987 for a review). Several 

studies of nonlethal disturbance have revealed that polychaetes regenerating damaged or lost body parts 

have lower fecundity and increased time to maturity than their undamaged counterparts (Hill and Grassle, 

1981; Hill et al., 1982; Zajac, 1985). 

Nonlethal disturbance could also affect the frequency of spawning of animals and plants. Barry (1989) 

suggested that disturbance might affect seasonal spawning patterns of Phragmatopoma californica

because lethal disturbance generally breaks tubes, facilitating a known spawning response. Young 

(1945) observed increased spawning frequency with increased stress on the byssal threads of the mussel 

Mytilus edulis. 

Spawning in individuals of detached mounds of P. lapidosa rolling around in the surf zone has been 

observed (J. Pawlik, University of North Carolina, Wilmington, NC, pers. comm.) Additionally, stress 

caused by changes in water pressure, desiccation, temperature, wave action, and salinity have been 

implicated in numerous studies of spawning and/or gametogenic synchrony of populations of marine 

invertebrates (see Giese and Kanatani, 1987 for a review). Differences in disturbance between habitats 

could affect fecundity differences between intertidal and subtidal individuals. Disturbance events that 

result in mortality can be very important in affecting the community of sessile organisms in a given area. 

On one end of the spectrum, high intensity and/or frequent disturbances such as wave activity can result 

in either no individuals in an area or very low numbers and diversities of organisms. In such cases, only 

the most morphologically adapted organisms such as limpets or chitons may be present. Moderate levels 

of disturbance may serve to mediate the effects of competition and result in the highest diversity of 

organisms (Levin and Paine, 1974). Such frequent disturbances may reduce the effectiveness of some 

opportunistic species from utilizing all available substrate space. The availability of such available space 

can then be colonized by other species if they can tolerate the level of disturbance in the area. In 

locations with lower intensity or less frequent disturbances, there is a tendency for lower diversity 

because a competitive dominant occupies most of the available substrate. During these competitive 

interactions, the size of substrate spaces created by disturbance is also important in terms of the type of 

organisms that occupy the space, and the time that it takes for the space to become fully utilized (see 

Sousa, 2001 for a review). Generally, larger “patches” of substrate space tend to be occupied more by 

sexually (versus asexually) reproducing species. These patches often take longer to become fully 

re-colonized. However, the time of colonization by sexual reproducers depends on how often these 

species spawn and the number of larvae they produce. In contrast, smaller patches tend to be colonized 

more quickly by asexually reproducing species (tunicates, corals, macroalgae) if they are present on the 

fringes of the patch. 
Section 3 7-15 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

7.5 OBLIGATE AND FACULTATIVE HABITAT USE 

Certain species such as P. lapidosa may be limited to NHB (some in IHB) because of the need for 

substrate to settle and on the turbulent hydrodynamic regime that aids tube building and feeding that is 

primarily found in these depths. Most other invertebrate species can be found on NHB, IHB, and OHB, 

although their proportions likely vary. A number of those invertebrates that take shelter within or eat 

worm rock may be considered obligate to P. lapidosa, but further empirical evidence is required. 

Hardbottom substrata provide attachment sites for most species of macroalgae. Many of these species 

probably have an obligate association with shallow hardbottom for shelter and position-maintenance. 

Many algal species also have adaptive chemical defenses against herbivores (Table 7.2). 

The species list for the entire NHB ichthyofaunal assemblage shows substantial similarities with many 

shallow coral reef species assemblages. Differences between NHB fish assemblages and reef 

assemblages in OHB depth ranges often involve the higher relative proportion of early life history stages 

on NHB structures, a juvenile-dominated assemblage. This suggests potentially important roles for 

differing ontogenetic species in shallow hardbottom systems (Werner and Gilliam, 1984; Livingston, 

1988), a useful concept for some applications involving the spatial ecology of species whose behaviors, 

habitats, and functional roles can change several times post-settlement. While NHB may be considered 

facultative habitat for juvenile green turtles because turtles can use alternate systems for growth (e.g., the 

Indian River Lagoon), it raises the question of what is the most essential or effective habitat for this life 

stage. The promotion of the potentially debilitating disease fibropapillomatosis (or FP) has been 

attributed to environmental cofactors present within lagoon/estuarine systems. The occurrence of FP is 

substantially higher in estuarine environments, making the Indian River Lagoon, overall, a less-than-ideal 

water quality habitat for turtles. Tracking data from several sources indicate that most sub-adult green 

turtles (approximately 70 cm SCL; D. Bagley, in prep.) relocate and move into seagrass beds, where they 

forage. Some important nutritional aspects that are components of marine turtle developmental phases 

may drive their selection of certain habitats. Little is known about hawksbill turtles in NHB habitats; 

however, their unique diet of sponges will limit their foraging range to more tropical regions in areas 

where NHB provides suitable substrate for sponge settlement. 

When newly settled and juvenile life stages are considered, questions of habitat dependence (an obligate 

attribute) and habitat opportunism (a facultative attribute) often focus on habitats used at settlement or 

associated life stages. Tests of dependence require consistent evidence that at least one life stage is 

restricted only to the habitat in question. For example, in the Dry Tortugas area of the Florida Keys, gray 

snapper settle in grass habitats in strictly euhaline waters and use a variety of other high-salinity habitats 

through maturity (Starck, 1970). Gray snapper can possibly migrate to Florida Bay from the Dry Tortugas, 

but because all demersal size classes occur in the euhaline, Dry Tortugas area, estuary dependence as a 

species paradigm is potentially excluded (Lindeman et al., 2000). Experimental manipulations could be of 

value in further examining the alternatives. 

For NHB-associated fishes, existing information can exclude many motile species as having an obligate 

association with shallow hardbottom. However, there are species such as the striped croaker, and some 

goby and blenny species, that can have very high associations with shallow hardbottom (and shallow 

artificial reefs). In these species, experimental studies could possibly parse the degree of habitat 

dependency. Importantly, obligate associations with specific habitats (and foods) are relatively 

uncommon for many motile, shallow fish species anyway; individual marine organisms still have an 

immediate need for the structure or food a facultative habitat provides. Therefore many habitats, though 

used facultatively, still have measurable and potentially essential value as aggregators of the shelter and 

food resources that the early life stages of many species require. 
Section 3 7-16 

and Mitigation Alternatives

Functional Attributes of Nearshore Hardbottom Table 7.2. Macroalgal species found on Florida’s Atlantic nearshore hardbottom habitat with known secondary metabolites 

defense against herbivores. Table constructed primarily from data in Paul and Alstyne (1992) and Diaz et al. (2

Species Asparagopsis (Falkenbergia hillebrandii) Dictyota indica Laurencia papillosa Avrainvillea sp. Dictyota pinnatifida Laurencia poiteaui Bryocladia cuspidata Dictyota sp. Laurencia sp. Bryopsis pennata Dictyota volubilis Lobophora (Pocockiella variegata

Bryopsis plumosa Galaxaura cylindrica Lyngbya sp. Bryopsis sp. Galaxaura marginata Neomeris annulata Bryothamnion seaforthii Galaxaura rugosa Neomeris sp. Bryothamnion sp. Galaxaura sp. Padina gymnospora Bryothamnion triquetrum Gelidiella sp. Padina profunda Caulerpa brachypus Gelidium americanum Padina sanctae-crucis Caulerpa cupressoides Gelidium crinale Padina vickersiae Caulerpa fastigiata Gelidium pusillum Padina sp. Caulerpa isthmocladium Gelidium sp. Penicillus dumelosus Caulerpa mexicana Grateloupia filicina Penicillus sp.Caulerpa microphysa Halimeda discoidea Polysiphonia binneyi* Caulerpa peltata Halimeda incrassata Polysiphonia denudata* Caulerpa prolifera Halimeda sp. Polysiphonia macrocarpa* Caulerpa racemosa Halimeda tuna Polysiphonia sp.*Caulerpa sertularoides Halymenia agardhii Polysiphonia sphaerocarpa* Caulerpa sertularoides farlowii Halymenia floresia Polysiphonia subtilissima* Caulerpa sp. Halymenia floridana Sargassum cymosum Caulerpa taxifolia Halymenia sp. Sargassum filipendula Chaetomorpha aerea Hincksia (Giffordia conifera) Sargassum filipendula var. montaChaetomorpha brachygona Hincksia (Giffordia duchassaigniana) Sargassum fluitans Chaetomorpha media Hincksia (Giffordia mitchelliae) Sargassum natans Chaetomorpha sp. Hincksia (Giffordia rallsiae) Sargassum polyceratium Codium decorticatum Hypnea cervicornis** Sargassum sp.Codium interextum Hypnea cornuta** Stypopodium zonale Codium isthmocladum Hypnea musciformis** Udotea conglutinate Codium repens Hypnea sp.** Udotea flabellum Codium sp. Hypnea spinella** Udotea sp.Codium taylori Jania adherens Ulva (Enteromorpha chaetomorpCryptonemia crenulata Jania capillacea Ulva (Enteromorpha compressa)Cryptonemia luxurians Jania rubens Ulva (Enteromorpha flexuosa parDictyota bartayresii Jania sp. Ulva (Enteromorpha flexuosa)* Dictyota cervicornis Laurencia corallopsis Ulva (Enteromorpha prolifera)* Dictyota dentata Laurencia gemmifera Ulva (Enteromorpha sp.)* Dictyota dichotoma Laurencia intricate Ulva lactuca* Dictyota dichotoma v. menstrualis Laurencia microcladia Ulva sp.*Dictyota divaricata Laurencia obtuse * Inducible defenses that operate quickly, from seconds to minutes. ** Defense under question. 
Section 3 7-17 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

7.6 POPULATION CONNECTIVITY AND MAINTENANCE 

For sessile invertebrates, there appears to be a trend towards limited dispersal of larvae or juveniles 

along the east Florida coast, but empirical information is limited. Of those species that reproduce 

sexually and asexually, many seem to propagate more via asexual reproduction. The scleractinian 

Acropora cervicornis is suggested to primarily reproduce when storm activity breaks its branches and 

individual pieces attach to the hardbottom (www.nmfs.noaa.gov/pr/species/invertebrates/staghorn.html). 

Recent molecular research supports this idea, as there appears to be restricted gene flow among 

Caribbean populations (Vollmer and Palumbi, 2007). Similar storm-induced asexual reproduction may 

influence reproduction in some sponges (Pione spp. and possibly Aplysina spp.) (Schönberg, 2002; 

M. Maldonado, Centros de Estudios Avanzados (CSIC), Girona, Spain, pers. comm.). 

In terms of sexually reproducing invertebrates, there is a tendency towards shorter planktonic periods, 

and larval behaviors that work in concert with local currents, which may restrict dispersal of some of these 

species (McCarthy, 2001; Vollmer and Palumbi, 2007). For example, P. lapidosa larvae can be ready to 

settle 2 weeks after fertilization (Eckelbarger, 1976; Pawlik and Mense, 1994). They likely position 

themselves deeper within the water column, which may allow for local retention (McCarthy, 2001; 

McCarthy et al., 2002). Florida populations genetically compared to those in the Caribbean appear 

different, suggesting limited genetic exchange (Drake et al., 2007). Finally, the ubiquitous shallow water 

scleractinian Siderastrea radians broods its young, likely resulting in lower dispersal of offspring from 

parent populations (Soong, 1993). 

A tremendous amount of new review, empirical, and modeling information is coming to bear on 

long-vexing issues of population connectivity. Considerable information has amassed regarding a variety 

of biotic and abiotic factors that contribute to retention in larvae. Retention on levels that challenge 

long-held views regarding long distance dispersal is now becoming commonplace (Jones et al., 1999; 

Cowen et al., 2000, 2006; Sponaugle et al., 2002; Paris et al., 2005; Cowen et al., 2006; Jones et al., 

2008). Though many differences exist, there are similarities between mainland east Florida shallow 

hardbottom habitats and rocky intertidal habitats of the U.S. Pacific coast at some scales of examination. 

Gunderson et al. (2008) proposed several broad categories of dispersal based on work with rocky 

intertidal Pacific fauna that have utility in examining potential dispersal and connectivity patterns in 

mainland east Florida. Their categories included 1) highly open connectivity among patchy populations 

and broad dispersal; 2) mesoscale dispersal structured by physics and resulting in several 

metapopulation sub-structures; 3) diffusive dispersal with stepping-stone metapopulations; and 

4) non-dispersing networks of closed populations. 

Efforts are underway to examine life history and empirical information on prominent invertebrate and fish 

species and functional groupings in the present study in order to assign select species or families of 

primary groups to appropriate connectivity categories. Nesting fish species with short larval periods 

(damselfishes), as well as species with limited pelagic larval periods (e.g., grunts, drums), show attributes 

of populations that could currently be placed in either the second or third dispersal category of Gunderson 

et al. (2008), through mesoscale details of east Florida differ. In terms of many taxa, these connectivity 

patterns may change markedly above and below the deflection of the Florida Current. Future research on 

these issues could be highly informative. 
Section 3 8-1 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

Chapter 8

Mitigating Shallow Hardbottom Impacts

An application of this synthesis report will be in the development of approaches to supplement current 

efforts to optimize the mitigation of unavoidable impacts. In this chapter, we examine fundamental theory 

and practice, and how mitigation is currently applied in nearshore waters of mainland east Florida. We 

also address Issue 4 (Chapter 1, Section 1.1), which is the question: if NHB has specific properties and 

functions that cannot be compensated for by existing OHB habitat, what are the best mitigation 

alternatives for the loss of NHB? 

To investigate the background issues and Issue 4, we used biological information from prior 

assemblage-specific chapters in Section 2 to examine preliminary scenarios of what happens to 

organisms during and subsequent to a dredge and fill burial at the fill site and how those stressors and 

effects may or may not cascade into local impacts at organismal and population scales. We also 

examine information regarding latitudinal differences in mitigation planning and implementation. It 

appears that only with such information aiding the front end of mitigation can the amounts and types of 

measurable compensation for lost services be fully evaluated. 

8.1 STRESSORS AND EFFECTS 

Recent methodological developments within the field of environmental assessment may aid evaluations 

of issues related to mitigation. Use of stressor and response characterizations within comparative 

ecological risk assessment frameworks is tractable. Full-experimental studies are often done in 

ecotoxicology, but synoptic, heuristic studies at scales that can slide from organismal to local population 

biology levels are feasible if the structured quantities of information assembled throughout Sections 2

and 3 are used. Whichever specific approaches are used, considerations of cumulative effects are 

necessary in comprehensive stressor and effect characterizations (Vestal and Rieser, 1995; Lindeman, 

1997b). 

8.1.1 Ecological Assessment Frameworks 

We used simultaneous evaluations of stressors, effects, and their potential co-occurrence with flexible 

protocols to help ensure comprehensive evaluations of effects (U.S. Environmental Protection Agency 

[USEPA], 1992; Harwell et al., 1995). This framework for assessing environmental effects of human 

activities was formalized by the USEPA in 1992 as comparative ecological risk assessment (CERA), and 

a number of approaches have developed. These approaches can be applied in iterative manners and by 

use of group-based protocols. 

Prior to examining mitigation issues in detail, we assembled a preliminary CERA to identify some of the 

effects for which a mitigation reef might have to compensate. We simultaneously evaluated information 

on the specific stressors introduced by an anthropogenic event, the biological responses of the organisms 

and life stages in question, and the ultimate effects at individual and population levels. Subsequently, 

information on stressors and effects was integrated based on spatial and temporal patterns of 

co-occurrence for prominent species groups. Due to the inherent complexity of characterizing stressors, 

effects, and their co-occurrence, full-scale ecological risk assessments are large projects, typically 

requiring a team of experts. Mechanics for executing a full CERA are detailed in the above references. 

8.1.2 Stressors, Responses, and Effects 

At least three fundamental categories of stressors can be identified in beach dredge and fill projects: 

turbidity, sedimentation (burial), and direct mechanical impacts. In the case of dredge and fill projects of 

mainland east Florida, sediments are excavated from areas offshore, fill is pumped into the supratidal and 

intertidal zones, a sand dike is built, and the fill is bulldozed into a broad beach. Most involve the 
Section 3 8-2 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

excavation and dumping of from 500,000 to 1,500,00 yd3 of sediments per project. Follow-up projects are 

usually incorporated into the original engineering design at 5- to 10-year intervals. Excavation typically 

occurs at one or more mid-shelf "borrow sites" that have been identified through geotechnical surveys as 

possessing beach-compatible sediments. Diverse spatial and temporal issues involving turbidity and 

sedimentation exist, including the abrasive effects of near-bottom sediment movements (V. Kosmynin, 

Bureau of Coastal Systems, FDEP, Tallahassee, FL, pers. comm.). The ratio of fines (silts and clays) to 

larger sediments is one of many multi-scale issues of particular environmental interest. 

In the present study, species of representative fishes, invertebrates, and algae were considered in terms 

of differential life stage-specific responses and effects (e.g., the ontogenetic species of Livingston, 1980, 

1988). It is not possible to characterize stressor effects and responses of every life stage of key species 

in association with dredge burial of their shelter and food resources. Considerable evidence suggests 

that survivorship of early life stages is a primary determinant of ultimate adult population sizes of reef 

organisms (Sale, 1991 and many other papers on marine invertebrates and fishes). Therefore, early 

demersal life stages, numerically dominant for many fish and invertebrate species (Sections 2 and 3), 

were emphasized in evaluations of potential responses. 

Table 8.1 shows multiple potential responses by physiological and behavioral systems (Olla et al., 1974; 

modified from Eisler, 1979). Many response categories can be associated with stressors arising from a 

natural or anthropogenic habitat burial event at short or extended temporal scales (Cech et al., 1998). 

Fish and invertebrate organ systems exist typically in states of relative homeostasis. Disturbance events 

create or amplify existing stressors. Organ- and behavior-scale systems can be challenged in discrete 

manners that, in combination, can rapidly degrade the fitness of cohorts of early life stages. 

Table 8.1. Representative response categories in early stages of marine invertebrates or fishes when 

challenged by stressors from habitat burial (Modified from: Eisler, 1974; Olla et al., 1974). 

Response Category Potential Response 

Sensory Capacities 

Phototaxis 

Geotaxis 

Chemotaxis 

Chemoreception 

Temperature preference 

Tactile inhibition 

Lateral line sensitivity

Rhythmic Activities Reproductive cycle 

Daily activity cycle

Motor Activities 

Avoidance 

Attraction 

Substrate association 

Equilibrium 

Swimming preference 

Spontaneous locomotor activities

Physiological Responses 

Respiration 

Feeding 

Pigmentation development 

Morphological development

Motivation and Learning Phenomena Feeding 

Conditioned avoidance response 

Migration Behaviors Subdaily feeding migrations 

Ontogenetic habitat shifts

Intraspecific Behaviors Visual attraction 

Aggregation and schooling

Interspecific Behaviors Aggression and territoriality 

Predation vulnerability
Section 3 8-3 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

In terms of the spatial distribution of stressors from dredge and fill activities, three overlapping cross-shelf 

strata are applicable: nearshore (0 to 4 m depth), intermediate (4 to 6 m), and shallow offshore (6 to 

10 m). The mechanical conditions under which many beach engineering operations occur can preclude 

direct assessment. For example, visual assessment of direct dumping of dredge fill on hardbottom reefs 

is not possible, often for weeks post-project, due to elevated turbidities at and adjacent to the project site. 

This situation also applies to direct assessment of the interface of the dredge head and substrate during 

excavation of fill offshore and habitat scale effects of fill burial inshore. 

Although effects of elevated turbidity on the health of organisms are poorly known in many marine 

environments (Rogers, 1990; Wilber and Clarke, 2001), some information is available. Offshore and 

inshore, the introduction of unusually high concentrations of silts and clays may exceed the tolerance 

thresholds of organisms in both direct (death) and indirect (lowered growth or lowered egg production) 

manners. A relatively short period of dredge-induced turbidity caused an abrupt decrease in growth in 

two species of hard corals and may have had long term impacts on coral reproduction, including the 

important Montastraea annularis (Telesnicki and Goldberg, 1995). This species is found on shallow 

hardbottom of Broward County and also is an important component of mid-shelf reef structure. Telesnicki 

and Goldberg (1995) also demonstrated negative effects of sediment loads on hard corals at turbidity 

levels below the maximum allowed by the State during dredging. Well-adapted invertebrates such as 

sabellariid worms that encrust shallow rocks can undergo partial or complete mortality in burial exceeding 

3 days (Sloan and Irlandi, 2007) and by the elimination of attachment substrata for future cohorts of 

larvae. 

In addition to characterizing effects and consequences at organismal and population scales, 

discriminating natural background variation from stressor-induced effects is also important across the 

shelf. Natural stressors can be significant in both winter and summer seasons. Long-term estimates of 

average turbidity values under natural conditions are lacking. In systems with substantial natural 

background disturbances, e.g., NHB of the northern project area, impact questions ideally should 

consider what additional stress loads beyond the natural background disturbance regime can generate 

measurable declines in primary metrics of NHB condition. On the other hand, perhaps the additional 

anthropogenic stress load is negligible with the pre-existing natural disturbance load. The problem is that 

it is not only a matter of “quantity” – days of high wind/wave energy or the frequency of natural vs. 

anthropogenic burial of NHB, but, in addition, “quality” – the durability of offshore vs. shallow sediments 

that are being displaced by dredges and storms, respectively. This physical complexity is multiplied by 

the physiological stressor-effect relationships of key species (e.g., Table 8.1). Figure 8.1 provides a 

schematic overview of some of the key biological process (growth rates, fecundity, mortality) in terms of 

their response to dredging scenarios in addition to natural background turbidity fluctuations. Importantly, 

these biological processes are related so that sublethal, lowered growth in a cohort can potentially lower 

fecundity at a later age (Figure 8.1). As with most multivariate, non-linear systems, conclusions 

regarding impacts will often be driven by site-specific aspects of the area in question and the breadth of 

species and life history stages considered. 

Along with hurricanes in summer, natural high-turbidity conditions in southeast Florida also result from fall 

through late winter wind events, often associated with fronts. For many species of fishes, this is 

apparently a period of less significance to settlement stages. Spring through fall dredge burials of 

hardbottom eliminate the habitat that marine fish and invertebrate larvae utilize right before or during their 

peak periods of settlement to bottom structures. For motile species, the assumption that most or all 

fishes and invertebrates on NHB habitats can simply swim away is tenuous when applied to newly settled 

and early juvenile life stages in a dredge-induced sedimentation event (e.g., see tables in 

Sections 8.1.2.1 and 8.1.2.2). 
Section 3 8-4 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

Figure 8.1. Three scenarios of cross-shelf turbidity and sedimentation effects (direct and indirect) for 

systems subjected to A) no dredging, B) some dredging, and C) semi-continuous dredging 

(e.g., north-central Jupiter Island). These scenarios assume substantial natural turbidity from 

storm events and represent effects of fill in addition to background turbidity levels. 

Early stages of fishes and invertebrates appear to be adapted to sheltering in hardbottom microhabitat 

crevices of all types and locations during wind and wave events and may survive such events. However, 

this assumes that the dredge fill sediments are coarse enough to not constantly resuspend. Laboratory 

and field investigations suggest that native beach sediments will settle to the bottom faster and resist 

resuspension more effectively than the high amounts of fine skeletal sediments transported to the fill sites 

from dredging of high carbonate sediments native to low-energy environments offshore (Wanless and 

Maier, 2007). Intermediate or long term information on changes in nearshore water clarity and other 

important physiological variables for fishes, invertebrates and algae are not available over the last 

decades. 

In a natural wind event, water turbidity stressors are probably near the same scale as in a full dredge and 

fill project. However, most natural wind events do not result in the complete burial of all hard bottom 

structure. Organisms that are adapted to survive natural wind and wave disturbance events rely on 

shelter; dredging projects steadily eliminate all of the shelter, unlike most storm events. There is clear 

evidence that hurricanes and tropical storms can move sediments in such volume that entire NHB tracts 

are buried. The additive or multiplicative affects of smaller wind/wave events versus major storm or 

dredge and fill events of less frequency suggest hypothetical and logical possibilities (Figure 8.1) but little 

empirics. If a precautionary approach to ecosystem-based management is being applied (e.g., National 

Research Council, 1999 and dozens of journal articles on coastal management and fisheries science), 

the collective impacts of dredge and fill projects cannot be negated simply by assumptions that major 

knowledge gaps (i.e., large topics with little published research) mean the array of potential stressors are 

benign in the long term (Peterson and Bishop, 2005). In addition, the recreational values and ecosystem 

services provided by these habitats within an overall degraded coastal system are in no way free from 

continued degradation as well. 
Section 3 8-5 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

In addition to natural turbidity and sedimentation events discussed above, there are influential and 

potentially synergistic effects from the discharge of freshwater through south Florida’s flood control 

system that create major turbidity plumes near inlets. During the wet season (May through October), 

normal discharges are less than 50 m3

/sec, but following hurricanes can reach 400 m3/sec (Hu and Wan, 

2006). High volume discharges are most prevalent near the St. Lucie Inlet in Martin County, because the 

St. Lucie River is the seaward discharge point for water released from Lake Okeechobee. Smaller 

volumes of freshwater are discharged from other inlets including Port Everglades in Broward County, 

Lake Worth and Jupiter Inlets in Palm Beach County, Fort Pierce Inlet in St. Lucie County, and Sebastian 

Inlet in Indian River County. In addition to influencing light penetration, turbidity, and salinity, freshwater 

discharges can carry contaminants, harmful algae, and excess nutrients. 

8.1.2.1 Invertebrates 

Stressors associated with sedimentation, turbidity, and mechanical impacts can affect co-occurring 

invertebrates on scales ranging from individual physiology through, potentially, local population structure 

(Table 8.2). Increased turbidity associated with dredging may cause sublethal impacts to sessile 

invertebrates that may include withdrawal into crevices or body cavities, and possibly termination of 

feeding. Similarly, motile invertebrates such as small crabs and shrimp may withdraw into crevices and 

be functionally prohibited from feeding because of not being able to seek food. With an increase in the 

scale of dredging, most of these organisms (regardless of the ability to move) would perish, or at the very 

least be severely physically and/or physiologically stressed. In the later case, such stress would likely 

reduce survival and reproductive ability (Table 8.2). Not only does this high mortality affect population 

numbers, it also likely affects trophic interactions (Nelson, 1989). 

Table 8.2. Invertebrate predictors of responses to dredge and fill burial: 1) organismal responses, and 

2) assemblage-scale responses. Scenarios are for the fill site only. 

1) Organismal Responses 

Scale of Effect Sessile Mobile on Bottom Highly Mobile 

Behavioral responses 

(min to hr) 

- Withdraw in crevices or 

within body cavity 

- Terminate feeding 

- May withdraw in crevices 

- Terminate feeding - Flee 

Physiological responses 

- Asphyxiation/death 

- Gill filament damage 

- Filter feeding shutdown 

- Other organ damage 

- Asphyxiation/death 

- Gill filament damage 

- Other organ damage 

- Predation in flight 

Affected invertebrate classes and 

life stages 

(All = all life stages) 

Demospongiae, All 

Turbellaria, All 

Anthozoa, All 

Polychaeta, All 

Hydrozoa, All 

Stenolaemata, All 

Anopla, All 

Gymnolaemata, All 

Enopla, All 

Gastropoda, All 

Bivalvia, All 

Crionoidea, All 

Polyplacophora, All 

Ascidiacea, All 

Archaeogastropoda, All 

Entoprocta, All 

Echinoidea, All 

Gastropoda, All 

Crustacea, All 

Merostomata, All 

Stelleroidea, All 

Holothuroidea, All 

Squid spp. (Teuthida) 

Behavioral responses 

(>1 day to 1 week) No survival Very low survival probability Attempt to disperse to other 

hardbottom habitat 

Flight access to other hardbottom Impossible Rarely Site-dependent 

Predicted local population mortality High, 95% to 100% 

High, 90% to 100% 

Sublethal impacts on any 

survivors 

Unknown 

Population connectivity Highly variable Variable 
Table 8.2. (Continued). 

Section 3 8-6 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

2) Assemblage Responses and Potential Local Population Implications 

Sessile Mobile on Bottom Highly Mobile 

Without mitigation Will cease to exist Will not exist or with greatly 

reduced numbers 

Will flee to other structural 

habitats, fate unknown 

With resource mitigation 

Mitigation for Shelter

In large part depends on recruitment capability of invertebrates. The recruitment of sessile shelter-enhancing invertebrates 

will significantly affect survival of motile invertebrate species. There will be very few older motile adults that will migrate into 

the affected area. 

Mitigation for Food Resources

If they can settle, most suspension feeding invertebrates should not be adversely affected. However, the slow recovery of 

crevice dwelling species (small crabs, shrimp, worms) may affect prey availability for many motile invertebrate predators and 

fishes. 

Water Quality

Degradation in water quality via immediate and chronic turbidity issues may affect fitness and/or survival of suspension 

feeders. It may also affect predator-prey interactions. 

Food web and community-scale trophic impacts 

Before

Community is highly diverse, consisting of a mix of opportunistic and rare species. Food web consists of suspension 

feeders, herbivores, omnivores, and various levels of carnivores. 

After

Low diverse community that consists of more opportunistic species and even fewer less abundant species such as corals 

and sponges. The reduction of shelter-enhancing sessile species will reduce the number of smaller carnivore trophic levels 

and is likely to affect higher trophic levels within the community. 

The loss of the high biomass of invertebrates (crabs, shrimp, worms, gastropods, bivalves) found within 

and near shelter-enhancing invertebrates (i.e., Phragmatopoma worms) and/or macroalgae can 

significantly modify in the nearshore food web. Before a dredging event, the community can be diverse, 

consisting of a mix of opportunistic and uncommon species. In this condition, the food web consists of 

suspension feeders, herbivores, omnivores, and various levels of carnivores. However, after a dredging 

event that results in high mortality and habitat elimination, there is likely to be a less diverse community 

consisting of more opportunistic species and fewer uncommon species such as corals. Reduced shelter 

enhancement (via fewer sessile species) will reduce the number of smaller carnivore trophic levels, with 

possible effects to the higher trophic levels within the community. 

Mitigation efforts to restore hardbottom habitats in large part depend on recruitment capability of 

invertebrates. The recruitment of sessile shelter-enhancing invertebrates will significantly affect 

recruitment and survival of motile invertebrate species. There will be very few older motile adults that will 

migrate into the area of impact. If they settle, most suspension-feeding invertebrates should not be 

adversely affected. However, the slow recovery of crevice-dwelling species (small crabs, shrimp, worms) 

may affect prey availability for many motile invertebrate predators and some fishes. Severe degradation 

in water quality (most likely via chronic turbidity) may affect fitness and/or survival of suspension feeders. 

It may also affect predator-prey interactions because of reductions in visibility. Significantly, mitigation 

planning should also consider that worm rock structures are most abundant in intertidal and subtidal NHB 

and substantially less adapted for IHB and OHB depths in east Florida (Chapter 4). 

8.1.2.2 Fishes 

The diversity of potential effects at the individual level is high. For example, seven differing sensory 

system variables and at least five other variables associated with motor activity and water-column 

positioning can be affected by dredge burials of resident fish habitats (Table 8.1). Effects can be direct or 

indirect and operate over the short-term, long-term, or both (Table 8.3). Indirect effects at the individual 

level can cumulatively translate into population-level effects or not. Impacts months or years after the 

initial impact are difficult to measure (Cech et al., 1998). 
Section 3 8-7 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

Table 8.3. Predictors of organism-scale responses to dredge and fill burial in fishes. Scenarios are for fill 

site effects only. 

Scale of Effect High Site Association Intermediate Site Association Fully Mobile 

Behavioral responses 

(min to hr) 

- Withdraw in crevices or 

within body cavity 

- Terminate feeding 

- May withdraw in crevices 

- Terminate feeding 

- Flee 

- Flee 

- Associate with sediment 

plume and feeding 

opportunities? 

Physiological responses 

- Asphyxiation/death 

- Gill filament damage 

- Other organ damage 

- Asphyxiation/death 

- Gill filament damage 

- Other organ damage 

- Predation in flight 

Affected fish families and 

life stages 

(All = all life stages) 

Pomacentridae, All 

Gobiidae, All 

Muraenidae, All 

Blenniidae, All 

Apogonidae, All 

Gobiesocidae, All 

Ophichthidae, All 

Pempheridae, NS, J 

Haemulidae, NS 

Sciaenidaea, NS 

Lutjanidae, NS 

Chaetodontidae, NS 

Pomacanthidae, NS 

Residents

Lutjanidae, J 

Scaridae, J, A 

Labridae, J, A 

Serranidae, J 

Sciaenidae, J 

Acanthuridae, J 

Balistidae, J 

Pomacanthidae, J, A 

Transients

Scorpaenidae, J, A 

Kyphosidae, J 

Chaetodontidae, A, J 

Pempheridae, A 

Lutjanidae, A 

Scaridae, A 

Scombridae, J, A 

Labridae, A 

Sciaenidae, A 

Acanthuridae, A 

Balistidae, J 

Haemulidae, A 

Carangidae, A, J 

Kyphosidae, J, A 

Sparidae, A, J 

Behavioral responses 

(>1 day to 1 week) No survival Very low survival probability Attempt to disperse to other 

hardbottom habitat 

Flight access to other 

hardbottom Impossible Very site-dependent Site-dependent 

Predicted local population 

mortality High, 95% to 100% 

High, 90% to100%. Mortality 

due to shelter or food resource 

gaps during flight periods of 

>1-2 days. Sublethal impacts 

on any survivors 

Low, but taxa-specific. Some 

species able to relocate with 

relative ease, others no 

Population connectivity 

Nesting species with low 

larval durations are 

common 

Variable Variable 

NS = Newly settled; J = Juvenile; A = Adult. 

Many responses will show some species specificity, and within species, substantial ontogenetic variation. 

The cryptic hardbottom resident fishes and herbivorous damselfishes (including adults) and the earliest 

life history stages of the other fish subassemblage types are among the most susceptible to direct 

mortality (Table 8.3). Consideration of effects on fishes at these scales, and their effects on higher 

organizational scales, are generally absent from the literature on shallow hardbottom and beach dredging 

in Florida. As the majority of motile fishes using nearshore areas of southeast Florida are juveniles or 

younger, and life stages of many conspecifics using mid-shelf areas are typically older, the greatest direct 

and indirect impacts may occur at nearshore fill sites and during flight responses; however, impacts at 

offshore borrow sites also remain underexamined, with occasional exceptions (Wilber and Stern, 1992). 

Many of the effects examined here may be pulsed through time on very patchy scales. Interactions 

among short- and long-term effects may also occur, adding to the complexity in evaluating stressors and 

responses. Examination of cumulative effects of both direct and indirect stressors is still often 

unattempted, yet of importance to the full understanding of anthropogenic effects. It is useful to 

discriminate among lethal and sublethal effects, which in turn can produce differing responses based on 

the time-scale evaluated. For example, identical organ systems can undergo sublethal or lethal effects 

(e.g., respiratory stress vs. respiratory trauma). These variations, in turn, lead to differing effects at 

greater exposure durations. Examples can be developed for each combination of stressor intensity and 

duration. 
Section 3 8-8 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

8.1.2.3 Algae 

Though over 200 species of algae are recorded from NHB, the diversity of potential effects may not be as 

high as in other organisms. Motility and other life history attributes are not as variable, and since almost 

all species rest on or attach to substrate, mortality is high across all divisions of algae (Table 8.4). 

Preliminary examinations of what effects such stressors can generate at population levels suggest that 

long term and more meso-scale spatial frameworks may be required to fully detail dredge and fill effects 

on algal populations and trophically-associated invertebrates and fishes (Figure 8.1). Potential long term 

reductions in water clarity can also affect food webs and require further examination to understand effects 

on growth rates, predation rates, and reproductive potential for diverse taxa. 

Table 8.4. Predictors of organism- and population-scale responses to dredge and fill burial in 

macroalgae. Scenarios are for fill site effects only. 

Scale of Effect High Site Association Intermediate Site Association Fully Mobile 

Behavioral responses N/A N/A N/A 

Physiological responses Mortality loss of photosynthesis 

and respiration Loss of primary production 

Affected taxa and life stages 

All Divisions 

Rhodophyta – SR, F, T 

Phaeophyta – SR, F, T 

Chlorophyta – SR, F, T 

Cyanobacteria 

Fragmentation/Clonal: 

Sargassum spp. 

Caulerpa spp. 

Spore fragmentation: 

Sargassum spp. 

Behavioral responses N/A N/A N/A 

Flight access to other 

hardbottom N/A N/A N/A 

Predicted mortality High, 75% to 100% Highly variable, 0% to 75%. 

Sublethal impacts present Low-none 

Population connectivity Quasi-closed, non-dispersing 

systems 

Diffusive and mesoscale 

dispersal Broad dispersal 

F = Fragment or clonal reproduction; SR = Spore recruitment; T = Turf species. 

N/A = Not available. 

8.1.2.4 Turtles 

NHB appears to provide intermediate life stage habitat for juvenile marine turtles, in particular green, 

loggerhead, and hawksbill turtles whose residency is approximated to be 2 to 5 years. Turtles may 

remain in the same area during this period or they may “migrate” with water temperature changes. The 

magnitude of site fidelity is unknown at this time, but tag and recapture data as well as tracking data 

indicate that turtles display some site specificity, even as juveniles (Ehrhart et al,, 1996; Bressette et al., 

1998; Kubis, 2003; Makowski et al., 2006b). Table 8.5 reflects preliminary efforts to assess the response 

of turtles to fill impacts given their mobility and potential responses. 

Poor water quality impacts food resources for most marine turtle species (i.e., molluscs and crustaceans 

for loggerheads, sponges for hawksbills, macroalgae for green turtles). Mitigation should include 

monitoring water quality before, after, and during dredge burial and mitigation construction. Water from 

land-use runoff sources should receive proper treatment before reaching the shoreline; otherwise, 

unhealthy and deleterious conditions may occur on both short- and long-term time-scales. Degraded 

water quality has been suggested as a potential environmental co-factor for FP disease found in a 

relatively large number of juvenile green turtle populations worldwide (Herbst and Kline, 1995). The 

disease manifests predominantly as tumors on the eyes and fleshy parts of the body. FP can impair 

swimming and foraging abilities and make an animal more susceptible to predation and/or starvation or 

death. Internal tumors are typically lethal. 
Section 3 8-9 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

Table 8.5. Predictors of organism- and population-scale responses to dredge and fill burial for marine 

turtles. Scenarios are for fill site effects only. 

Scale of Effect High Site Association Intermediate Site Association Fully Mobile 

Behavioral responses 

- Flee to deeper water 

- Withdraw under ledges 

- Loss of food resources 

- May withdraw under ledges, 

or flee in random walk 

response, or flee in optimal 

foraging response. 

- Immediately flee 

Physiological responses - Increase foraging effort 

- Increased predator exposure 

- Increased predator exposure 

- Slower growth rate 

- Stress 

- Increased predation 

- Slower growth rate 

- Stress 

- Higher carbohydrate intake 

necessary for energy to 

migrate 

Affected taxa and life stages 

Green turtle - J 

Hawksbill - J, S 

Loggerhead - S 

Disperse until new structure 

present 

Green turtle - J 

Hawksbill - J, S 

Loggerhead - S 

Loggerhead - S 

Behavioral responses see above see above see above 

Flight access to other 

hardbottom 

Green turtle 

Loggerhead turtle 

Hawksbill turtle 

Transient/migratory? 

Green turtle 

Loggerhead turtle 

Hawksbill turtle 

Green turtle 

Loggerhead turtle 

Hawksbill turtle 

Predicted mortality Unknown/undocumented Unknown/undocumented Unknown/undocumented 

Population connectivity 

Quasi-closed as juveniles. 

Juveniles appear to disperse at 

certain size-classes 

Juveniles appear to disperse at 

certain size-classes 

Broad dispersal (temporary 

condition) 

J = Juvenile; S = Subadult. 

8.2 MITIGATION OF NEARSHORE HARDBOTTOM BURIAL 

The process and practice of mitigation is similar to ecological restoration; the distinction between 

restoration and mitigation is that mitigation is invoked when the impact is known in advance, not 

accidental or natural as in restoration. Thus the relatively new field of restoration ecology provides much 

of the theoretical and applied direction for mitigation efforts (e.g., Race and Fonseca, 1996; Zedler, 2000; 

Falk et al., 2006; Miller and Hobbs, 2007). Most of the experience in attempting to restore habitats and 

ecosystems comes from terrestrial and wetland environments. The legislative framework for mitigation is 

also derived primarily from wetland contexts. Thus far, work in marine environments in general and NHB 

mitigation in particular is still in its infancy. 

As discussed in Chapter 1, adherence to particular ecological concepts can influence the interpretation of 

restoration efforts. In fact, restoration projects are themselves likely to yield new insights into ecological 

understanding, as many are quasi-experimental. Downes et al. (2002) stated “It has been said that an 

ultimate test of our understanding of an ecosystem is to create or repair a habitat and its function.” 

As discussed above, dredging projects designed to rebuild eroded beaches may bury expanses of NHB 

and increase turbidity, both acutely and chronically. Although NHB is naturally buried and uncovered 

seasonally and with the passage of storms, additional burial through dredging can amplify and extend the 

natural effects and loss of habitat within a stratum of the broader cross-shelf continuum. This is 

particularly true if burial affects time periods specific to the larval recruitment or movements of local fishes 

and invertebrates. 

For projects conducted under Section 404 of the Clean Water Act or the Rivers and Harbors Act of 1890, 

potential impacts are treated in a three-step sequence. The first part of the sequence seeks through the 

analysis of alternatives to avoid adverse environmental impacts; if adverse impacts cannot be avoided, 

then all efforts to minimize them must be taken; and in the event that minimization is not feasible and 

impacts are unavoidable, then compensatory mitigation is required. In most beach dredging projects 

where hardbottom is present in the project area, compensatory mitigation is required (USFWS, 2004). 
Section 3 8-10 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

Mitigation efforts should seek to ameliorate not only the effects of direct burial and immediate habitat loss 

but also turbidity, including chronic turbidity. Artificial reefs can compensate only for structural habitat 

loss; mitigating for chronic turbidity and low water clarity (transmissivity) is not accomplished by reef 

placement. In this chapter, the focus is on mitigation of direct habitat loss through burial by dredge fill. 

To date, deploying artificial reefs has been the preferred route for compensating for unavoidable impacts 

to NHB (USFWS, 2004). Artificial reefs have been used for fishery management and habitat 

enhancement for years (Seaman, 2000; Brickhill et al., 2005), but the use of reefs for mitigation or 

restoration has been a more recent phenomenon (Ambrose, 1994; Miller, 2002; Reed et al., 2006). The 

process usually involves pre-project evaluation of expected impacts to existing NHB by utilizing aerial 

photographs to quantitatively map the area of hardbottom to be impacted. Expected impacts are 

modeled by estimating where the seaward margin of fill will distribute along the shoreline, the equilibration 

point following placement of sand on the beach. Once the potentially impacted area is determined, a 

mitigation plan is prepared, consisting of a calculated mitigation ratio, an engineering analysis, and a 

monitoring plan. 

8.2.1 Artificial Reefs 

The approach used to offset the effect of habitat burial has been to construct artificial reefs near impacted 

sites. Because some fishes and invertebrates will colonize hard substrate through migration of adults or 

settlement of larvae from the plankton, simply placing a structure on the seafloor will attract certain 

elements of the local fauna. As mentioned above, artificial reefs have a long and complicated history in 

fishery and habitat management. In addition, artificial structures have been used extensively in Europe 

and other coastal areas in association with beach projects (Bacchiocchi and Airoldi, 2003; Airoldi et al., 

2005). 

Research on the applications of artificial reefs has been extensive (e.g., Seaman and Sprague, 1991; 

Seaman, 2000), with legitimate claims of success, typically for offshore fishing values. Artificial reefs can 

become ecological traps that subject colonizing organisms to fishery exploitation (Bohnsack, 1989) or 

high levels of predation (Hixon and Beets, 1993), and they can be sources or sinks within particular 

seascapes (Crowder et al., 2000). Success is often a function of what is measured, and consensus 

agreement on the most critical metrics for success often is not evaluated. 

Placement of reefs on sedimentary bottoms can, over time, alter sediment grain size distribution and 

organic matter accumulation around the reef, causing a corresponding change in associated infaunal 

assemblages (Davis et al., 1982; Ambrose and Anderson, 1990; Fabi et al., 2002). None of these 

drawbacks should preclude the use of artificial reefs as mitigation unless independent research from the 

proposed deployment site indicates that one or all would be influential. 

A commonly posed concern with the use of artificial reefs is the potential predation risk to newly settled 

organisms. It is assumed that predators are more abundant or more effective on artificial reefs than they 

are on natural reefs. It is important to note that the field studies that demonstrated high predation were 

conducted on small artificial reefs, where the presence of only a few small predatory fishes can exert 

considerable influence on the existing community (Hixon and Beets, 1993; Eklund, 1996; Leitao et al., 

2008). It is not clear how the results from studies of smaller isolated artificial reefs can be scaled to larger 

artificial reefs, but certainly increasing or decreasing the size or complexity of a reef could modify 

observed predation effects. McCarthy (2001) studied post-settlement mortality of sessile invertebrates in 

intertidal and subtidal hardbottom off Boynton Beach and found no predation effect for total number of 

species or for the majority of individual species. This suggests that sessile invertebrate mortality in 

intertidal and shallow subtidal hardbottom can be the result of sand scouring and wave activity in addition 

to capture by mobile predators such as crabs or fishes. 

Because current mitigation efforts have been exclusively centered on constructing artificial reefs in 

shallow water, an essential step in the process is to evaluate engineering stability. Reefs placed in 

shallow depths to achieve like-kind mitigation will be subject to high-energy waves and dynamic sand 

movements. Most reefs have been built from concrete or quarried limestone boulders (Figure 8.2). 

These materials are subjected to stability tests that consider a range of wave energy impinging on 
Section 3 8-11 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

proposed structures. Stability test results generally preclude deploying limestone boulders or concrete 

slabs in water depths less than 4 m. These depths also mark the shallow operating depths of work 

barges and attending tugboats. 

Figure 8.2. Artificial mitigation reef and substantial algal and hydroid growth, 6 m depth, Juno Beach, 

Palm Beach County, Florida. Photo by D. Snyder.

An additional consideration is sediment overburden within proposed reef footprints. If the sediment 

overburden exceeds the maximum dimension of a reef unit, the entire reef unit could settle under the 

sand. Therefore, prior to selecting sites, considerable effort must be expended to measure sediment 

thickness (e.g., Deysher et al., 1998). 

Because of the stability and logistic issues stated above, it is apparent that NHB cannot be mitigated for 

in like-kind fashion, given present technology and budgets. The solution has been to build reefs in 

deeper water where they are persistent and not subject to the burial and erosion that characterizes the 

0 to 4 m depths (Figure 8.2). Reefs built of limestone boulders are higher relief with different structural 

features than much of the naturally occurring NHB. In addition to stability and logistic concerns, spatial 

arrangement of artificial reefs over the seafloor needs to be evaluated from an ecological perspective 

(e.g., Jordan et al., 2005; Lindberg et al., 2006). These factors must be considered when monitoring and 

evaluating the efficacy of mitigation reefs. Alternative modular designs are also being developed 

(Figure 8.3; Continental Shelf Associates, Inc. et al., 2006). 

In assessing restoration efforts in terrestrial assemblages by reference sites, White and Walker (1997) 

classified the basic elements of the reference information based on proximity in space and time. Their 

classification, revised for NHB, is presented in Table 8.1. The actual impact site provides the current 

ecological snapshot of conditions of the impact site, but it is only a snapshot and does not give an 

accurate picture of processes. For example, in coastal areas the natural condition may have long ago 

been obscured by anthropogenic effects, leading to the question “what is natural?”(Sapp, 1999; Jackson, 

2002). Temporal data from the impact site are valuable, particularly if a before-after-control-impact 

(BACI) approach is to be used in assessing the restoration (Osenberg et al., 2006). Data such as these 

would be short term, for example during the year prior to a proposed dredge and fill project. Longer term 

(decadal) patterns will be more elusive, as White and Walker (1997) point out that temporal data do not 

always allow us to directly associate physical and biological factors related to disturbance regimes. 

Contemporary information from different sites will provide insight on how existing processes affect 

species assemblage attributes and dynamics. Unfortunately, in most NHB situations the primary spatial 

factor is one of water depth. 
Section 3 8-12 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

Figure 8.3. Prototype mitigation reef modules made from articulated mat with integral coquina rock 

surface. Each block is about 30 inches x 30 inches x 12 inches (Continental Shelf 

Associates, Inc. et al., 2006). Photo by D. Snyder.

If like-kind mitigation is desired, then managers and permit applicants will have to accept that material will 

be lost (buried) under the sand. To some extent, this mimics the natural situation. Given the expense of 

constructing reefs, it is unsettling to place one in very shallow water only to have it disappear beneath a 

moving sand bar. However, during other seasons the reef material will likely uncover. Potentially more 

problematic would be public safety issues in very shallow waters as well as potential movement of reef 

materials onto the beach with time. 

Fauna and flora that live in the intertidal sections of the NHB have to be adapted to survive through 

changes in a number of environmental factors. These factors not only change with season but also with 

state of the tide. They include seasonal and daily changes in water/air temperature, salinity, and 

desiccation. Additionally, organisms within this depth zone are exposed to some of the highest wave 

energy that occurs when seasonal storms or hurricanes affect the area. 

Restoration of buried worm reefs could be better effected by using existing knowledge on behavioral 

preferences of P. lapidosa and other sessile invertebrates during settlement. For instance, it has been 

well documented that P. lapidosa repeatedly “test” substrates before settling on the preferred one 

(Eckelbarger, 1976; Pawlik, 1988). The recruitment of P. lapidosa to deployed substrates is variable but 

commonly higher at locations closer to the bottom and/or with moderate turbulence (D. McCarthy, 

Jacksonville University, Jacksonville, FL, pers. observ.). Behavioral preferences for moderate amounts of 

current have been quantified in laboratory studies with P. californica (Pawlik et al., 1991; Pawlik and 

Butman, 1993). Consequently, assuming adequate larval supply, there are likely to be unique 

combinations of hydrodynamics and substrate features that will enhance recruitment of invertebrates. A 

description of those features would allow coastal managers to modify artificial reefs to include those 

preferred features and thereby maximize larval settlement and formation of “natural” habitats. 

McCarthy and Holloway-Adkins (2007) investigated whether the P. lapidosa will recruit on deployed sets 

of settlement plates in ~4 m water depth off Brevard County, Florida, an area that has little significant 

subtidal NHB. They also investigated whether recruitment varies with height above the seafloor bottom, 

orientation, or chemical coating of the settlement plates. They deployed three 100 cm x 100 cm x 70 cm 

ultraviolet-stabilized polyethylene propagule and larval measurement (PALM) boxes off Satellite Beach, 

Florida (28°09.637’ N, 80°34.951’ W) in a water depth of 4.6 m. Fifteen 10 cm x 10 cm x 1.5 cm 
Section 3 8-13 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

limestone settlement plates were vertically attached on the north, east, west, and south faces of each 

PALM box. The PALM boxes were deployed for two time periods. The first deployment occurred on 

24 May 2006, and plates were recovered on 8 July 2006. The second deployment also took place on 

8 July 2006, with boxes being recovered on 5 May 2007. 

They found that P. lapidosa recruitment varied considerably for the two time periods sampled. During the 

May to July 2006 time period, P. lapidosa was the dominant encrusting organism, recruiting in fairly high 

yet variable levels. Most P. lapidosa worms encountered on the settlement plates were alive and in a 

range of sizes indicating multiple cohorts of recruitment. Additionally, encrusting species such as 

bryozoans, hydrozoans, ascidians, and barnacles were also frequently encountered on the settlement 

plates. During the July 2006 to May 2007 time period, the PALM boxes were partially buried with 

sediment and experienced high mortality of recruits of various species of encrusting organisms. 

P. lapidosa recruitment was very low, with most of the observed recruits on the plates being bivalves and 

barnacles. For both time periods, there was a trend that P. lapidosa recruitment was lowest on plates at 

the rows furthest from the seafloor. Otherwise, P. lapidosa recruitment occurred equally on plates 

regardless of chemical treatment and plate orientation. In conclusion, recruitment of P. lapidosa did occur 

to artificially deployed structures off the Brevard County coast in mean water depths of approximately 

4.6 m. 

The variation in recruitment between sampling periods may be because of 1) differences in time the 

plates were deployed, and/or 2) natural seasonal fluctuations in larval availability. The general lack of 

significance among the experimental treatment means are probably a result of local hydrodynamic and/or 

turbidity conditions that consistently and dramatically fluctuate. These fluctuations likely are continually 

creating favorable conditions for the settlement of P. lapidosa larvae, regardless of the effect of plate 

orientation, height, and chemical treatment. Finally, it should be cautioned that P. lapidosa may recruit at 

these latitudes but may not survive at these depths because water temperatures likely reach cold 

temperature limits for this species (Eckelbarger, 1976; D. McCarthy, Jacksonville University, Jacksonville, 

FL, pers. comm.). 

The use of artificial reef habitat by marine turtles is largely undocumented in the peer-reviewed literature. 

Jones et al. (2004) observed juvenile green turtles resting among artificial reef rocks deployed in 

approximately 4 to 7 m water depths in Boca Raton, Florida. Garrido (2007) observed hawksbill turtles on 

shallow artificial reefs in West Palm Beach, Florida. Juvenile green and adult loggerhead turtles were 

observed on the nearshore artificial reef during annual surveys between 2006 and 2008 (Coastal 

Eco-Group Inc., 2008, 2009). Properly designed artificial reefs with ledges and crevices may provide 

adequate shelter for resting marine turtles and provide adequate substrate to support the foraging habits 

of hawksbill and green turtles. However, juveniles mostly forage on shallow, narrow reefs in intertidal and 

shallow subtidal depths, and the placement of mitigation reefs in deeper waters may create competition. 

These aspects remain unstudied, to date. 

Mitigation reefs provide hard substratum for the recruitment of macroalgae and invertebrate food 

resources for fish, turtles, and other invertebrates. Mature macroalgae have been successfully 

transplanted on artificial reef surfaces (Ohno et al., 1990). Holloway-Adkins and McCarthy (2007) 

conducted experiments to test macroalgal recruitment on subtidally deployed structures off Brevard 

County where hardbottom is absent. The experiment involved examining levels of recruitment on 

different substrate types that might be used in artificial reef construction in 4.6 m water depth. While 

characteristics of limestone (light coloration, porosity, and texture) make it ideal substrata for macroalgae 

(Dawes, 1998), no significant differences in macroalgal recruitment on experimentally tested settlement 

plates were found (Holloway-Adkins and McCarthy, 2007). Six of 15 red algae and three of five green 

algae documented by Continental Shelf Associates, Inc. (2005a) on adjacent NHB recruited on the 

settlement plates. Eleven macroalgae and seven invertebrate species found in the foraging samples of 

green turtles recruited on the deployed settlement plates (Holloway-Adkins and McCarthy, 2007). 

Monitoring of mitigative artificial reefs in Broward County revealed overall macroalgal species richness 

was as high, or sometimes higher, as the natural NHB reefs during post-construction monitoring periods 

(Prekel et al., 2007). Species richness of green turtle-preferred macroalgae after 36 months was also 
Section 3 8-14 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

similar to what was found on natural NHB. However, higher quantities of macroalgae were documented 

on the natural reefs than on the artificial reefs (Prekel et al., 2007). 

The deployment of mitigation reefs prior to dredge and fill projects could, ideally, provide a more timely 

compensation of functional loss through the continuous existence of hardbottom structure that would 

provide shelter and recruitment habitat during and after project construction. The timing of mitigation 

deployment, both seasonally and pre-construction, could potentially influence initial recruitment success. 

Deployment of mitigation reef structures 6 months to 1 year prior to project initiation could allow for more 

adequate dispersal of macroalgal spores and fragments, as well as invertebrate larvae from existing 

hardbottom. There have been relatively few attempts to mitigate losses of NHB along the east Florida 

coast until recently. In 2004, the USFWS (2004) reviewed permit applications from southeastern Florida 

(Indian River to Dade Counties) for projects where compensatory mitigation was recommended or 

required to offset impacts to hardbottom habitats. They selected 26 projects from USACE regulatory 

division permits and planning division civil works projects. This review revealed that 217 acres of 

hardbottom was expected to be impacted. Approximately 113 acres of artificial or natural materials were 

to be deployed as replacement habitat; this total included 43 acres completed and 70 acres for projects 

not yet completed (USFWS, 2004). The impacts in these examples were caused by filling, sedimentation, 

dredging for beach nourishment, and port expansion (Table 8.6). The anticipated impacts of pending 

projects were also related to beach nourishment and port expansion. 

Table 8.6. Summary of compensatory mitigation for completed projects in southeast Florida between 

1985 and 2004 (Adapted from: U.S. Fish and Wildlife Service, 2004). 

Project Type Name/Corps’ Florida 

County 

Avoid and 

Minimize 

Reduced 

Impacts 

(acres) 

Recommended 

Acres 

Required 

Acres 

Constructed 

Acres Location 

Type 

(in/out of 

kind) 

Material 

(Acreage) 

Beach Renourishment 

Indian River County, 

Sectors 1&2 (200091872) 

Indian 

River Yes 7.2 10.5 5.2 Offsite In Limestone

Fort Pierce Shore Protection 

(Federal project) St. Lucie Yes N/A 5.0 5.0 5.0 Onsite In Limestone

Martin County 

Renourishment (199501665) Martin Yes .38 N/A 1.4 5.0 Onsite Out Concrete 

Jupiter/Carlin Park Shore 

Protection (199900902) 

Palm 

Beach Yes N/A 4.0 4.0 4.0 Offsite In and Out

Limestone 

(3.25) 

Concrete 

(0.75) 

Jupiter Inlet Sand Bypassing Palm 

Beach No N/A 2.32 N/A 1.16 Offsite In Limestone

Juno Beach Shore 

Protection (1997066559) 

Palm 

Beach Yes 

Reduced 

fill and 

length 

4.47 4.47 2.22 Onsite and 

Offsite In Limestone

Phipps Park Shore 

Protection 

Palm 

Beach Yes N/A 3.1 N/A N/A Onsite In Limestone

Ocean Ridge Shore 

Protection (199301576) 

Palm 

Beach Yes 5.9 4.55 4.55 4.55 Onsite and Offsite In and Out

Limestone 

(2.1) 

Concrete 

(2.0) 

North Boca Raton Shore 

Protection 

Palm 

Beach Yes 0.2 N/A 0.16 0.16 Offsite In 

Limestone 

(0.07) 

Modules 

(0.09) 

South Boca Raton Shore 

Protection (199401196) 

Palm 

Beach Yes N/A 2.38 2.38 3.0 Offsite In Limestone

Broward County Shore 

Protection (19990554) Broward Yes 22 13.5 7.6 8.9 Onsite In Limestone

Bal Harbor BEC&HP 

(Federal project) 

MiamiDade N/A N/A 0.8 N/A 0.48 Offsite In 

Limestone 

(0.26) 

Modules 

(0.22) 

Sunny Isles BEC&HP 

(Federal project) 

MiamiDade Yes N/A 3.13 3.13 0.33 Onsite and 

Offsite In 

Limestone 

(0.19) 

Modules 

(0.14) 

63rd Street BEC&HP 

(Federal project) 

MiamiDade Yes N/A 0.08 0.08 0 Offsite In Limestone
Table 8.6. (Continued). 

Section 3 8-15 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

Project Type Name/Corps’ Florida 

County 

Avoid and 

Minimize 

Reduced 

Impacts 

(acres) 

Recommended 

Acres 

Required 

Acres 

Constructed 

Acres Location 

Type 

(in/out of 

kind) 

Material 

(Acreage) 

Surfside and Miami Beach 

BEC&HP (Federal project) 

MiamiDade Yes 0.005 0.07 0.07 0.07 Offsite In Limestone

Total -- -- 13.69 36.58 30.44 25.97 -- -- -- 

Navigation Improvements 

Fort Pierce Deepening St. Lucie Yes 2.25 15.0 4.0 4.0 Offsite Out Concrete 

4.0 

Hillsboro Inlet Navigation 

Improvement (199301995) Broward Yes 1.25 0.8 0.8 0 Onsite In Limestone0 

Miami Harbor Deepening MiamiDade No 0 N/A 13.5 13.5 Onsite In 

Limestone 

N/A 

Modules 

0.6 

Total -- -- 3.5 15.8 18.3 17.5 -- -- -- 

Combined Totals 

(Beach/Nav) -- -- 17.19 52.38 48.74 43.47 -- -- -- 

N/A = Not available. 

For 18 completed projects, USFWS (2004) identified approximately 103 acres of direct impact to 

hardbottom habitat. Of this, 15 acres were related to port channel expansion, 49 acres were covered 

during beach nourishment projects, 25 acres were due to long-term sedimentation from pipeline leaks or 

rupture, and 14 acres were related to pipeline or anchor cable placement. An additional 10 acres of 

indirect impact to NHB was projected from short-term turbidity and sedimentation at Ocean Ridge and 

Indian River County beach nourishment projects. 

Another 10 proposed projects were expected to directly impact about 114 acres of hardbottom habitat. 

Eighty-three acres are expected to be impacted from proposed port expansion, 18 acres are expected to 

be impacted from dredge filling, 12 acres are expected to be impacted by the deployment of liquid natural 

gas pipelines, and 1 acre is expected to be impacted from sedimentation (USFWS, 2004). Since the 

2004 USFWS report was released, there have been changes to some of the projected projects. For 

example, Brevard County Mid-Reach will impact approximately 3 acres of hard bottom, not 22 acres 

(Continental Shelf Associates, Inc. et al., 2006). For 15 of the 18 completed projects, compensatory 

mitigation was recommended, and the total acreage was 69 of the 103 acres directly impacted by these 

projects (Table 8.6). For the proposed projects, mitigation was recommended for 6 of the 10 proposed 

projects. These six projects will account for 82 acres of the 100 acres expected to be directly impacted. 

8.2.2 Monitoring 

Monitoring of mitigation performance or success is an important part of the entire process. Great sums of 

money are spent with current mitigation programs, and it is imperative to gain an understanding of 

successful applications to better guide future attempts. Monitoring mitigation projects is similar in 

approach to environmental impact monitoring, but there are important differences. The two monitoring 

types are similar in that most situations do not lend themselves to elegant experimental designs. Impact 

or mitigation locations are rarely replicated, and suitable control areas are often hard to find. Currently, 

there are several issues to consider when establishing a particular monitoring of restoration or mitigation 

program: 

• Setting goals; 

• Framing hypotheses; and 

• Study design. 

Establishing the goals of the monitoring program should be the first priority once the impact acreage has 

been scaled to corresponding mitigation acreage. Intuitively, the goals for NHB would be to restore the 

functional attributes of the impacted habitat to the level of an adjacent natural reference area. In reviews 

of past restoration projects, particularly in wetland and terrestrial contexts, clear statements of the project 
Section 3 8-16 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

goals often were not included (Race and Fonseca, 1996; Hackney, 2000; Redmond, 2000). Setting goals 

will influence most of the other elements of the process. For example, if the goal is to restore functional 

aspects of the impacted assemblage, then the focus should be on reestablishing functional groups or 

clusters of focal species. Osenberg et al. (2006) distinguished between endpoint and effect-size based 

goals, pointing out that both approaches have merits. Endpoints provide a basis for comparison and 

efficacy, whereas the effect size is necessary to understand how well the mitigation performed. 

Again, the conceptual foundation of ecology comes into play, and interpretation of various concepts such 

as community structure, function, and succession will be important in setting and evaluating goals 

(Palmer et al., 1997). Application of various community ecology theories in restoration efforts will depend 

on species present and characteristics of the natural environment. In the nearshore environment where 

assemblages would be less biologically predictable and driven by chance colonization and recruitment, 

conventional succession theory may be of little use. 

Framing scientific and statistical hypotheses in accordance with standard practice in ecology 

(falsificationist approach) relative to restoration projects presents a logical paradox (McDonald and 

Erickson, 1994; Underwood, 1996; Downes et al., 2002). In the conventional approach, the researcher 

frames a null hypothesis, such as: there is no difference in mean species richness between the control 

and impact sites. Data are collected and evaluated using a statistical test for the hypothesis that is 

rejected if an arbitrarily selected p-value for the test is reached. As Underwood (1990) pointed out: 

“It is sometimes considered appropriate to phrase the statistical definition of recovery as some measure in 

the disturbed area not being significantly different from that in control areas. This has been fossilized in 

legislation in some aspects of environmental management (USEPA, 1985). This is inappropriate because 

sloppy, imprecise sampling leads to declaring recovery is complete when it is not. If the confidence 

interval(s) around the mean measure in the disturbed area or in the control areas (or both) is (are) large, 

they will not differ significantly in any test—even if the mean values are nothing like each other.” 

Statistical methods exist that remedy these logical issues; these methods are called bioequivalence by 

McDonald and Erickson (1994). With this approach, the researcher defines, in advance, a minimum 

measure. For example, 90% of the fish species richness (or 80% of the biomass of epibiota) found in 

control areas would represent recovery in the mitigated habitat (e.g., McDonald and Erickson, 1994; Cole 

and McBride, 2004). This quantitatively defined endpoint must be exceeded before the habitat is 

considered restored. Thus, with this method the null hypothesis is that the mitigation site is not equivalent 

to the control site. Bioequivalence gets around the logical problem, and the formal hypothesis tested is 

ratio of variables between the restored and reference sites. Following this approach, inadequate 

sampling will cause the test to retain the null hypothesis of no equivalence, even if the mean measure 

does exceed the criterion for recovery. Furthermore, the burden of proof is now with the responsible 

party. 

In the past, most deployments were made with engineering considerations mentioned above and have 

not lent themselves to elegant experimental designs. Researchers are forced to use approaches that 

deal with lack of replication, arbitrary placement of materials, and other violations of conventional 

experimental treatments. For this reason, the various forms of BACI designs are most appropriate 

(Osenberg and Schmitt, 1996; Stewart-Oaten and Bence, 2001; Osenberg et al., 2006). In addition, 

control site selection is a problem in many areas where NHB is impacted. The ideal control site is not a 

pristine and unaffected site against which to compare the mitigation reef. Control sites are those selected 

to represent the habitat that is being mitigated for, including any current anthropogenic influences. They 

are not randomly selected from a population of such sites. Equivalence tests described above can be 

used with BACI designs (Downes et al., 2002). 
Section 3 8-17 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

A variation of the BACI design is to restore only part of an impacted area and compare the restored site 

(artificial reef) to the “unrestored control sites” as well as a natural control site over time (Downes et al., 

2002). This will allow researchers to judge the direction of change, as well as the relative degree of 

change. Downes et al. (2002) indicated that three distinct treatments are needed: 

1. degraded location(s) that need restoration; 

2. similar degraded site(s) that will not be restored; and 

3. site(s) that represent the target state. 

A critical element of study designs is identifying response variables and using those variables to evaluate 

success. McCoy and Mushinsky (2002) described success as “an acceptably small difference between 

the structure and/or function of a restored system and that of a reference system.” When convergence of 

community structure is the goal in evaluating restoration success for artificial reef deployments in NHB 

mitigation, this definition could be incorporated into an equivalency statement. As mentioned above and 

by others (Underwood, 1990; Downes et al., 2002), the challenge is determining the response variable(s) 

and setting the level of equivalency. 

8.3 CONCLUSIONS 

8.3.1 Regional and Organism-specific Issues 

• Mitigation of NHB impacts by using artificial reefs can often provide adequate replacement of lost 

structural habitat and therefore some ecological functions. 

• Not all reef designs and deployments will be successful in all situations; site-specific factors must be 

considered. Different approaches and designs will be required for different locations in southeast 

Florida and east-central Florida. 

• For worm reef areas, ensuring successful local larval recruitment events is essential. To this end, 

project designers need to be sure that sediment size range is suitable to incoming recruits. 

• For worm reef areas, a measure of success is not only the presence of worm reef but duration: it 

should persist through 1 year with new recruits being encountered in the second year. More 

importantly, there should be some of the dominant crustaceans encountered within it 

(e.g., Pachycheles monilifer [porcellanid crab], Pachygraphsus transverses [grapsid crab], Menippe 

nodifrons [xanthid crab], and Synalphus fritzmuelleri [snapping shrimp]). 

• In deeper or more southern areas of the project region, the presence of specific sponge and 

anthozoan species can be used to judge success of artificial mitigation reefs. 

• In shallow mitigation to the mid- and north sections of the project region, reef deployment should 

minimize impact to the yearly recruitment cycles of the habitat engineering worm reef. 

• In terms of Issue 4: Can NHB functions be replaced with artificial reefs in IHB or OHB depths? 

Probably not in large part for IHB and less for OHB, due to the depth limits on the habitat engineering 

worm species P. lapidosa, and the associated assemblage of many dozens of species that 

P. lapidosa structures support in areas from Brevard County to mid-Palm Beach County. 

• Artificial reef mitigation does not appear to remove or provide compensatory mitigation for the direct 

and indirect physiological and potential population scale impacts of turbidity and sediment 

resuspension. 

• If possible, materials should be used for artificial reef construction that support algae and worm rock 

to the north, and more sponges and corals to the south. 

• Over 275 species of algae associate with NHB, and most are subject to high mortality on sites that 

are filled. Mitigation reefs can provide new attachment sites for algae. Algal growth on mitigation 

reefs should correlate with water clarity. 

• Pre-construction deployment of mitigation reefs (6 months to 1 year prior) in regards to seasonal 

recruitment patterns of focal reef organisms (i.e., sabellariid worms, prominent fish species, 

macroalgae) may potentially accelerate functional compensation of habitat loss and should be a 

priority in project planning. 
Section 3 8-18 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

8.3.2 Monitoring Issues 

• A single approach for scaling mitigation is needed early in the project. Although competing models 

can be effective, using two different (non-complementary) approaches on the same project can lead 

to confusion. 

• Clear goals for mitigation should be established. 

• Comparisons of biotic attributes between natural and artificial reefs should be framed using 

equivalency approaches when feasible. 

• Study designs should be flexible and adaptive given the typical lack of natural controls or replication. 
Section 3 9-1 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

Chapter 9

Summary

Synthesis of available information on shallow hardbottom structure and distribution, organismal biology 

and assemblage scale drivers, and stressors and mitigation responses are presented within Sections 1, 

2, and 3, respectively. Although many research gaps are present, substantial amounts of information can 

be assembled relative to shallow hardbottom ecological functions in terms of latitudinal and depth 

variation in areas within the study region. We will summarize primary findings on assemblages of algae, 

invertebrates, fishes, and turtles. We will then summarize information on ecological functions of shallow 

hardbottom, stressors and responses, and mitigation alternatives. 

In the context of the three sections we will also consider the four original issues: 1) What are specific 

ecological functions of NHB?, 2) What is happening to the coastal ecosystem (broader than NHB) when 

this NHB is lost due to nourishment?, 3) Is OHB able to compensate for the loss of NHB?, and 4) If the 

NHB habitat has specific properties and functions that cannot be compensated for by existing OHB 

habitat, what are the best mitigation alternatives for the loss of natural NHB? Issue 1 and portions of 

Issues 2 and 3 are addressed in sections below on algal, invertebrate, fish, and turtle assemblages. 

Additional components of Issues 2, 3, and 4 are subsequently addressed in sections below on ecological 

functions. Issue 4 is addressed in the section on mitigation with support from other sections. A paucity of 

depth-specific data exists for a majority of taxa in the region and continues to limit some summary 

statements about core differences among NHB, IHB, and OHB. 

9.1 ALGAL, INVERTEBRATE, FISH, AND TURTLE ASSEMBLAGES 

Shallow hardbottom habitats of southeast and east central Florida support a sometimes diverse, 

juvenile-dominated reef assemblage of invertebrates, fishes, and marine turtles. There are also many 

species of algae that can dominate cover. In total, at least 1,050 algal, invertebrate, and fish species are 

now recorded from NHB habitats of mainland east Florida. The physical environment is subject to wind 

and wave disturbance of sometimes high levels at intermediate temporal and spatial scales. Hardbottom 

patches occur in many areal sizes, frequencies, and depths and are the predominant shallow water 

habitat off mainland Florida’s beaches. These patches are the only natural structural habitat, outside of 

sand plain features, on the ocean side of all the inlets from Government Cut north to at least Port 

Canaveral; commonly considered habitats such as seagrass beds and mangroves do not occur outside of 

the inlets along this reach. Patchily distributed NHB, IHB, and OHB areas can aggregate many diverse 

organisms and can be of local significance within the nearshore shelf seascape. 

9.1.1 Algae 

The total number of recorded algal taxa, identified and unidentified, is approximately 340, including 

277 known species. Potentially dominant genera are Ceramium, Dictyota, Hypnea, Laurencia, Gelidium, 

Caulerpa, Jania, and Ulva, depending on subregions and other factors. While no one macroalgal species 

can be considered a keystone in the NHB, a handful of species appear to repeatedly surface in our 

examination of abundant and frequently utilized macroalgae. Halimeda spp. also aid in sand production. 

Sunlight converted into macroalgal tissue is directly consumed by as many as 20 genera of invertebrates, 

at least 14 genera of fishes, and the juvenile stage of the endangered green turtle, Chelonia mydas. 

Algae are a dominant driver of the food web and also contribute to shelter used by dozens of species of 

invertebrates and fishes. All algal species contribute to oxygen and nutrient production. In terms of 

functional form groups, jointed-calcareous algae are most responsible for sediment enrichment and sand 

building. 

The abundance and diversity of macroalgae on NHB vary substantially with changes in latitude from 

Cape Canaveral to Miami Beach. Proximity to relatively warm, oligotrophic Florida Current waters is a 
Section 3 9-2 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

primary factor influencing macroalgal species distribution on NHB. Several other factors that influence 

distribution include small and large scale hydrodynamics, different substrate types (e.g., worm reefs, 

limestone, coral reefs), water depth, light penetration (due to water clarity), sand scouring and burial, 

wave energy, fish and invertebrate grazing pressure, and the various life history strategies of different 

macroalgae (Lobban and Harrison, 1994). Some observations on the distribution of algae on NHB are 

1) overall biomass appears comparatively higher in northern NHB regions (i.e., Indian River and Brevard 

Counties), 2) species with more grazer-resistant properties (e.g., phaeophytes, calcareous and crustose 

rhodophytes) are more diverse and abundant in southern NHB regions, and 3) cyanobacteria are 

commonly encountered on all NHB. However, caution is advised in drawing conclusions from these 

observations since they have yet to be thoroughly and rigorously tested. A more thorough understanding 

of the functional role of macroalgae and cyanobacteria would include measuring: a) resilience under 

disturbance events (e.g., hurricanes), b) abundance and diversity at macro- and micro-level scales, 

c) temporal variability, and d) role of grazers in controlling productivity. 

9.1.2 Invertebrates 

Over 533 species of invertebrates have been identified along the east Florida coast. The diversity of 

sessile species is greatest for cnidarians (~21%), bryozoans (~29%) and sponges (~19%). Some sessile 

taxonomic groups may not be represented with such high diversity but can occur in very high biomass. A 

primary example is the sabellariid polychaete Phragmatopoma lapidosa, which can be very abundant 

along the mid- to north sections of the project area and is a habitat engineer that creates structure 

supporting high diversities of many other invertebrates. 

Sessile invertebrate communities change noticeably with latitude and depth. In intertidal hardbottom to 

the north, dominant invertebrates are sabellariid polychaetes, bryozoans, sea anemones, and tunicates, 

although the latter three groups generally do not occur in high biomass. Throughout intertidal areas, in 

addition to P. lapidosa, the hard coral Siderastrea spp., two species of zoanthids, and several species of 

solitary anemones can be encountered. Within the lower intertidal and shallow subtidal zones, the most 

conspicuous sponge is the rock-boring sponge Pione (formerly Cliona) lampa. In the more centrally 

located sections of the east Florida coast, there is generally an increase in abundance of P. lapidosa both 

intertidally and subtidally to water depths of approximately 4 m. Within Martin County, the St. Lucie Inlet 

Reef appears to be the northernmost extent of several of the reef-building corals that are important 

biodiversity contributors within the Caribbean region. The biomass of hard and soft corals and sponges, 

while variable and generally low, tends to increase to the south. 

The most diverse and generally abundant motile invertebrates are arthropods and polychaetes, which 

represent 47% and 28%, respectively, of the total number of motile invertebrate species. Over 

100 species of crustaceans are extremely abundant, especially on worm reef-dominated hardbottoms. 

These include crabs, stomatopods, shrimp, isopods, and amphipods. There are over 87 reported 

polychaete species that are likely to be very abundant, although more research is needed to confirm this. 

There are also fairly high numbers of gastropods, flat worms, ribbon worms, and echinoderms on these 

habitats. While less diverse, there generally are intermediate numbers of echinoderms such as sea 

urchins to the north of the coast. Some groups such as brittle stars are probably fairly diverse and 

abundant but have not been extensively studied. 

Primary ecological functional roles of invertebrates along the east Florida coast include 

1) shelter-enhancing organisms that increase local diversity of fishes and invertebrates, and 2) predators 

or prey in local food webs. Generally, the highest community biomasses along the mainland east Florida 

coast occur in hardbottom areas with higher abundances of sessile invertebrate species (some may be 

considered foundational or keystone contributors to the community) that enhance local shelter. Along the 

Florida coast, important shelter-enhancing taxonomic groups are hard and soft corals, sponges, 

tunicates, molluscs, barnacles, and polychaetes (i.e., P. lapidosa). However, their importance in this 

function may vary dramatically with depth and latitude. In terms of contribution to local food webs, 

important taxonomic groups are sponges, crabs, shrimp, polychaetes, echinoderms, crabs, and shrimp. 
Section 3 9-3 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

The loss of NHB would result in a significant reduction of a high biomass of invertebrates (sponges, 

corals, crabs, shrimp, worms, gastropods, bivalves) that could significantly change the nearshore food 

web. Before a dredging event, the nearshore community can be fairly diverse, consisting of a number of 

common and longer-lived (often uncommon) species. In this situation, the diversity contributes to a 

nearshore food web that consists of suspension feeders, herbivores, omnivores, and various levels of 

carnivores. However, after a dredging event occurs that results in high mortality and habitat elimination, 

there is likely to be a less diverse community consisting of more opportunistic species, and fewer rare 

species such as corals. Further, conditions where there is significant reduction in shelter enhancement 

(via fewer sessile species) in these areas could reduce the number of smaller carnivore trophic levels and 

have possible adverse effects on the higher trophic levels within the community. Natural recovery of 

hardbottom habitats would depend on both re-exposure of substrate as well as recruitment capability of 

sessile shelter-enhancing invertebrates which in turn likely affect associated motile invertebrate species. 

Although it requires further investigation, it appears that very few older adult motile invertebrates would 

migrate into the area of impact. Consequently, habitat recovery requires ample larval supply and 

recruitment of shelter-enhancing species to facilitate what could be a slow recovery of many associated 

motile invertebrates. The slow recovery of crevice-dwelling species (small crabs, shrimp, worms) may in 

turn affect prey availability for many motile invertebrate predators and some fishes. Further, severe 

degradation in water quality (most likely via chronic turbidity) may affect fitness and/or survival of 

suspension feeders. It may also affect predator-prey interactions because of reductions in visibility. 

It is unlikely that OHB is able to compensate for the loss of NHB, particularly in the northern areas of the 

east Florida coast. Mitigation efforts to restore hardbottom habitats likely depend in large part on 

recruitment capability of shelter-enhancing species such as P. lapidosa. Therefore, local hydrodynamics 

and the type, topography, and relief of materials must be considered for artificial reefs to maximize 

recruitment and subsequent recovery of the habitat. For instance, in the case of P. lapidosa, the 

abundance of this species decreases significantly in water deeper than 4 m. Even within the 0 to 4 m 

depth range, the highest abundances occur more towards the intertidal zone. Why this occurs may be 

related to improved feeding and reproduction that may occur in these more turbulent shallow waters. 

However, the choice to settle and live there depends entirely on factors that affect larval supply. It is 

likely (though requires further investigation) that settling P. lapidosa larvae may touch bottom at deeper 

waters but actively postpone metamorphosis and choose to settle when they encounter shallower waters 

(McCarthy, 2001). Consequently, if the goal is to mitigate for loss of worm reefs (and associated 

organisms), then structures deployed in the OHB would not be successful unless chemical cues, 

hydrodynamics, or other environmental factors could be created that accurately mimic the shallower 

waters and induce larvae to settle. Further, if settlement does occur at these depths, growing juveniles 

must survive and recruitment of associated species must occur to properly mitigate for lost habitat. 

9.1.3 Fishes 

Based on new and old information, over 257 species of fishes are now recorded from NHB habitats. The 

species list for the entire ichthyofaunal assemblage shows similarities with assemblages from many 

shallow coral reefs (Chapter 5). A wide range of spawning patterns is seen among the fishes of east 

Florida NHB. Nesting is undertaken in differing manners by species of fishes from quite different families. 

Several different families, including the site-attached blennies and gobies, also build nests in cryptic 

manners throughout NHB structure, often laterally or upside-down in NHB microhabitats. Though 

understudied, we estimate that over 30 fish species spawn on NHB. Most motile species do not spawn 

on NHB, though there are exceptions. In addition, some coastal pelagics may spawn on or near NHB. 

Although the assemblage is juvenile-dominated, a well-developed food web exists that includes most or 

all common reef trophic categories, including planktivores, herbivores, invertivores, and piscivores. The 

most prevalent trophic component among species is invertebrate feeders. Randall (1967) categorized the 

diversity of invertivores according to 1) sessile animal feeders (e.g., sergeant major feeding on 

Zooanthus); 2) “shelled” invertivores (e.g., most Halichoeres species of wrasses), and 3) generalized 

invertebrate predators. The latter group was by far the biggest and included eels, squirrelfishes, 

snappers, species of basslets (Serrranus), some drums (sciaenids), goatfishes (mullids), pompanos, 

wrasses, and some labrisomids, including the hairy blenny. Specialized invertebrate feeding can occur 
Section 3 9-4 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

as well (Chapter 5). When the full spectrum of ontogenetic dietary shifts is considered for most species, 

many are omnivorous (e.g., planktivory to invertivory to piscivory), from species as diverse as labrisomids 

and lutjanids. 

Are nearshore fish larvae spending substantial time in the water column near NHB before settling there? 

This is possible for some taxa, particularly clupeiforms, gerrieds, sparids and others, but currently cannot 

be confirmed. For many species in south Florida, transport of larvae to NHB or OHB settlement areas 

may occur via cross-shelf oceanographic transport mechanisms resulting from diverse meso-scale 

phenomena (Limouzy-Paris et al., 1997). Directed water column or bottom sampling as done in some 

other areas (Ruple, 1984; Jahn and Lavenberg, 1986) is needed to address this issue. 

Differences among ichthyofaunal assemblages are suggested between the southern and northern areas 

of mainland southeast and east central Florida in terms of the most abundant species and habitat 

structure (Chapter 5). Potential explanations for substantial differences in prominent species (sailors 

choice, black margate, hairy blenny) within a 360 km north-south gradient may include temporal 

recruitment variations, NHB structural differences between the two areas, differences in survey design, 

species reaching their biogeographic distributional limits between the northern and southern areas, and 

other factors (summaries of Chapters 5 and 7). 

Bellwood (1998) concluded that reef fish assemblages ultimately occur in association with too many 

non-coral reef structures and recommended the term fishes on coral reefs. However, Robertson 

(1998a,b) defined reef fishes as “species that live on consolidated substrata that form coral and inorganic 

reefs.” Inorganic reefs, including rocky shores, were described as: “... the inorganic bedrock provides 

large and small physical structure that fishes use as habitat features...”. All of the 10 consensus families 

of reef fishes in Bellwood (1998) are represented in association with NHB of east Florida. This evidence 

suggests there are few reasons to conclude that the fishes of NHB of mainland east Florida do not 

constitute a reef fish assemblage. 

9.1.4 Turtles 

Juvenile sea turtles of three species commonly associate with shallow hardbottom of mainland east 

Florida: the green (Chelonia mydas), loggerhead (Caretta caretta), and hawksbill (Eretmochelys 

imbricata) turtles. There are many interactions between turtles and an array of organisms that utilize 

shallow hardbottom. For example, hawksbill turtles fragment sponges when foraging, creating space and 

habitat for other organisms (Leon and Bjorndal, 2002) as well as increasing sponge growth through 

fragmentation. Juvenile stages of the endangered green turtle can associate with shallow hardbottom for 

years, feeding on macroalgae and using structure for shelter. And while green turtles that forage on 

seagrasses have been examined for their important role in the productivity and nutrient cycling of 

seagrass communities (Thayer et al., 1984; Aragones and Marsh, 2000; Moran and Bjorndal, 2007), the 

role of green turtles foraging on macroalgae in macroalgal communities has not been examined until now 

(K. Holloway-Adkins, ongoing research). Under conditions where substrate for the attachment and 

growth of macroalgae are limited and herbivory is high, grazers can potentially experience resource 

competition or partitioning. The mechanistic selection behavior under variable resource conditions is 

important to understand, especially when planning projects that potentially impact the plight of 

endangered species. 

Conditions over many NHB sites make studying marine turtle populations difficult, and only a handful of 

studies have been conducted. Juvenile green turtles are the most frequently encountered species on 

NHB. Genetic stock studies utilizing mitochondrial DNA analysis were conducted on a subset of juvenile 

green turtles captured in two long-term (>10 years) population studies on NHB: Bagley (2003) in Indian 

River County and Bass and Witzell (2000) in St. Lucie County. Results indicate that NHB juvenile green 

turtle populations in these areas were a mixed genetic stock with the same genetic haplotypes as those 

from an array of different nesting beaches in Costa Rica, Florida, Mexico, and Aves Island (Caribbean). 

Juvenile hawksbill turtles have only recently been studied and are rarely encountered north of Martin 

County. Genetic data from turtles captured on NHB, IHB, and OHB indicate they represent a genetic 

stock from various Mexico nesting beaches (L. Wood, Palm Beach Zoo, pers. comm.). 
Section 3 9-5 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

Data indicate that green and hawksbill turtles predominantly are the most frequently encountered species 

and the ones that assume relatively long-term residency on NHB. These data and observations suggest 

that the developmental life stages of juvenile green and hawksbill turtles in particular are intimately tied to 

the NHB habitat. The mean population size and size-class range suggest that the smaller mean size 

turtles (20 to 42 cm CL) are obligately tied to certain habitats within the NHB. 

9.2 ECOLOGICAL FUNCTIONS AND CONNECTIVITY 

Many species use NHB, IHB, and OHB as settlement and early juvenile habitats. The great majority of 

the species are recorded from coral reefs. For fishes, NHB assemblages are largely juvenile dominated 

and reef-species dominated. As the only natural habitat structure for 0.5 to 5 km of sand in any direction, 

shallow hardbottom structural features potentially serve a variety of ecosystem functions, including 

settlement and nursery areas, spawning sites (for over 100 species of invertebrates and fishes), feeding 

stations, and socioeconomic drivers (see Chapter 7). The ecosystem service roles of shallow 

hardbottom to local resident water users (fishers, divers, surfers), their families, and the local communities 

along mainland east Florida remain under examined. Conceptual ecological models can be a useful way 

to organize the bio-physical and socioeconomic complexity of shallow coastal systems (Ogden et al., 

2005). Empirical information on feeding relationships will allow construction of more resolved food webs, 

which should enhance the development of conceptual ecological models for shallow hardbottom areas. 

It is useful to consider the larger scale that these local populations are distributed across (in part, via 

planktonic larval stages). Outside of ahermatypic hardbottom reefs, nearshore marine areas of mainland 

east Florida show no natural, high relief, three-dimensional structures that support fish, algal, and 

macroinvertebrate assemblages. Energetically, hardbottom areas are nutrient-rich patches in among 

large sand plains. Ecological functions include shelter and feeding resource provisions to over 

1,050 recorded species. 

Currently, the majority of NHB habitat is within a narrow, 200- to 400-m strip eastward of a major north-tosouth reach of beaches from Melbourne to Miami and display a variety of structural forms. Notable 

exceptions with much more gradation of shallow hardbottom habitats across the shelf and into deeper 

areas occur in Broward and southern Indian River Counties. NHB composes less than 50% of the 

alongshore distance of any of the seven county shorelines in the region, where it is often separated by 

kilometers of flat nearshore sand expanses. Latitudinal abundance differences may be common for 

prominent species (Sections 2 and 3). 

For sessile invertebrates, there appears to be a trend towards limited dispersal of larvae or juveniles 

along the east Florida coast. Of those species that reproduce sexually and asexually, many seem to 

propagate more via asexual reproduction. Recent molecular research supports this idea, as there 

appears to be restricted gene flow among Caribbean populations. Similar storm-induced asexual 

reproduction may influence reproduction in some sponges (Pione spp. and possibly Aplysina spp.). 

Much new review, empirical, and modeling information is coming to bear on issues of population 

connectivity (Chapter 7). Considerable information involving similar reef organisms from other areas has 

been amassed regarding a variety of biotic and abiotic factors that contribute to retention in larvae (e.g., 

Cowen et al., 2006). Retention on levels that challenge assumptions of routine long distance dispersal is 

now becoming more commonplace. Though many differences exist, there are similarities among 

mainland east Florida shallow hardbottom habitats and rocky intertidal habitats of the U.S. Pacific coast at 

some scales of examination (Sections 2 and 3). 

9.3 MITIGATION 

Increasingly important and expensive questions involve the use of mitigation projects to offset impacts 

from large dredge and fill projects. Typically, these questions represent key steps in the finalization of 

permitting processes. Preliminary conclusions regarding the functional attributes of NHB and the 

application of that information to the evaluation of mitigation alternatives include the following: 
Section 3 9-6 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

• Mitigation of NHB impacts by utilizing artificial reefs can often provide adequate replacement of lost 

structural habitat and therefore some structure- and trophic-associated ecological functions. 

• Not all reef designs and deployments will be successful in all situations, and site-specific factors must 

be considered. Different approaches and designs will be required for different locations in southeast 

and east central Florida. 

• For worm reef areas, ensuring successful local larval recruitment events is essential. To this end, 

project designers need to be sure that the sediment size range is suitable to incoming recruits. 

• For worm reef areas, a measure of success is not only the presence of worm reef but duration: it 

should persist through 1 year, with new recruits encountered in the second year. More importantly, 

some of the dominant crustacean species should be encountered within the worm rock. 

• In terms of Issue 4, based on available information, NHB functions will not be fully replaced with 

artificial reefs in IHB or OHB water depths due to the depth limits on the habitat engineering worm 

species P. lapidosa and the associated assemblage of many dozens of species that P. lapidosa

structures support in areas from Brevard County to mid-Palm Beach County. 

• Artificial reef mitigation does not appear to remove or provide compensatory mitigation for the direct 

and indirect physiological and potential population-scale impacts of turbidity and sediment 

resuspension. 

• Approximately 340 algal taxa are associated with NHB, and most are subject to high mortality if a site 

is filled. Mitigation reefs can provide new attachment sites for algae. Algal growth on mitigation reefs 

should correlate in part with water clarity. The most abundant species on NHB should be considered 

in success criteria for artificial reef mitigation progress and monitored for persistence. 

• Invertebrate and fish grazers on artificial reefs placed in IHB or OHB waters may be subject to higher 

predation (especially in the more northern NHB regions where high-energy waves reduce the number 

of larger organisms in intertidal and shallow subtidal areas). Exclusion experiments on natural NHB 

vs. artificial mitigation reefs placed in deeper waters will reveal if mitigation reef productivity is 

reduced by increased grazer pressure. 

• Deployment of mitigation reefs at least 6 months to 1 year prior to project construction could 

accelerate colonization of algae and invertebrates, as well as provide alternate habitat for animals 

able to escape burial. 

• Placement of artificial reefs in IHB or OHB waters to mitigate for NHB dredge and burial projects 

should consider the possibility that where smaller size-class juvenile marine turtle populations exist, 

reefs placed in deeper waters may create habitat and resources suitable for larger turtles that could 

displace smaller individuals. Project planning should include pre- and post-monitoring criteria that 

include designing artificial reefs that provide adequate shelter for the existing size-class of juvenile 

turtle populations. Predation may also be increased on artificial reefs. 

• Monitoring plans for juvenile marine turtles should include documenting both size and species, either 

through observer training and/or the capture and measuring of individual animals. 

• Hypotheses in mitigation studies should be framed using equivalency approaches; this will also 

contribute to goal setting and success criteria. 

• Study designs should be flexible and adaptive, given the lack of natural controls or replication. 

9.4 RESEARCH NEEDS 

In terms of both ecological functions of shallow hardbottom and mitigating the loss of NHB, there are 

fundamental questions that require basic and reliable estimation of long-term patterns. Understanding 

effects beyond project boundaries at larger spatial scales is important to understanding project 

significance. Some areas of additional work include the following: 

• Latitudinal abundance differences in prominent species require more research to explain the 

north-to-south differences in assemblage pattern. 

• What are the most useful functional categories that can be evaluated from available information for 

shallow hardbottom habitat/organism assemblages? Manipulative experiments could be of particular 

use for some questions. 

• With evaluation, are such categories applicable to the shallow hardbottom system? 
Section 3 9-7 

Functional Attributes of Nearshore Hardbottom 

and Mitigation Alternatives 

• A food web model for shallow hardbottom is needed for multiple purposes. Enough information to 

build such a model from the bottom levels up is now accessible; however, empirical trophic studies 

are still badly needed. 

• A full conceptual ecological model (CEM) needs to be developed; this will be more efficient if a 

thorough food web draft is available. The information to build a larger CEM is now available for some 

components. 

• The intermediate or long term water quality patterns and effects on shallow hardbottom from turbidity 

and sedimentation events, long-shore and cross-shelf, that result from dredge and fill projects (and 

relationships to natural turbidity) require examination (Bishop and Peterson, 2005). Without this type 

of information, responses to Issue 2 regarding whole ecosystem effects when NHB is buried, or 

Issue 3 regarding OHB compensation roles, will be not be fully resolvable. 

• Experimental investigations of mitigation reefs are needed to examine extant questions about boulder 

reefs vs. modular reefs, depth-related issues, and performance of differing reef array designs. 
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Chapter 10

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Appendices 

Appendices
Appendices A-1 

Appendix A
Appendices A-2 Table A.1. Number of macroalgal species found during studies conducted on nearshore hardbottom habitat, by county. Da

study (region), species found, zone of sampling, method used to collect data (i.e., natural reef or artificial reef, b

monitoring, or foraging analysis, etc.). Color-coding indicates that similar study methods were used. County Region Species Count Intertidal Subtidal Inshore Offshore PCM - T Study Method 

Brevard coastline 24 √ √ Video, random point analysis SSatellite Beach 37 √ PALM boxes SNearshore reef, Brevard County 24 √ Lavage-percent composition 

Brevard Brevard County 22 

Vero Beach 109 √ √ Transect SSebastian 46 √ √ Lavage-percent composition 

28 √ Quadrat-percent composition 

Indian River Indian River 42 

Hutchinson Island 119 √ Trawl/benthic grab Se

- 

Fort Pierce 29 √ √ 

St. Lucie Fort Pierce-SC 51 

St. Lucie Inlet 41 √ Transect (drift algae included) Nearshore Mitigation Reef 9 √ √ (Post) Video, photo transect 

Martin Martin County 9 
Table A.1. (Continued). 

Appendices A-3 County Region Species Count Intertidal Subtidal Inshore Offshore PCM - T Study Method 

Boca Raton artificial reef 10 √ Quadrat-percent cover Boca Raton nearshore reef 23 √ Transect Palm Beach County 11 √ Lavage-percent composition 

N

Boca Raton 17 √ Grab samples Artificial reefs: (1) Juno Geogrid (2) Jupiter Cloth Reef Rock (3) Jupiter Concrete (4) Jupiter Shallow Concrete (5) Coral Cove Rock 12 √ √ (Post) Video, random point analysis Palm Beach County 26 North Jupiter 9 Jupiter Area 6 South Jupiter 6 Juno 8 Palm Beach MacArthur Beach 5 Broward County 10 √ Necropsy and lavage Fort Lauderdale √ √ √ Lauderdale by the Sea √ √ √ Pompano Beach √ √ √ John U. Lloyd State Park √ √ √ Dania √ √ √ Hollywood/Hallandale 27 (total) √ √ √ AGRRA protocol, in-situ quadrat analysis SPort Everglades- Broward/Miami-Dade County 100 √ √ Transect, BEAMR quadrat, video, random point analysis 

Broward Shoreline (Segments I, II, III) 6 √ √ Visual transect S
Appendices A-4 

Table A.2. Families, genera, and numbers of species within each genera of macroalgae and 

cyanobacteria documented in the studies conducted on nearshore hardbottom of the east 

Florida coast listed in Table A.1. A zero under “Number of Species” column indicates that 

algae were identified only to genus level. 

Family Genus Number of Species 

Chlorophyta 

Bryopsis 2 Caulerpaceae Caulerpa 12 

Chaetomorpha 3 

Cladophoraceae Cladaphora 6 

Rhizoclonium 2 

Codiaceae Codium 5 

Batophora 1 

Cymopolia 1 

Dasycladus 1 Dasycladaceae 

Neomeris 1 

Derbesiaceae Derbesia 1 

Halimedaceae Halimeda 6 

Dictyosphaeria 0 

Siphonocladaceae Siphonocladus 1 

Ventricaria 1 

Avrainvillea 2 

Udoteaceae Penicillus 1 

Udotea 6 

Ulvaceae Ulva 6 

Ulvellaceae Entocladia 1 

Rhodophyta 

Acanthaceae Acanthophora 2 

Acrochaetiaceae Acrochaetium 2 

Areschougiaceae Eucheuma 2 

Asparagaceae Asparagopsis 1 

Callithamnion 2 

Centroceras 1 

Ceramium 9 

Crouania 1 

Griffithsia 1 

Spermothamnion 3 

Spyridia 3 

Ceramiaceae 

Wrangelia 2 

Champiaceae Champia 1 

Compsopogonaceae Compsopogon 1 

Amphiroa 4 

Fosliella 2 

Jania 4 

Lithophyllum 0 

Corallinaceae 

Titanophora 1 

Corynomorphaceae Grateloupia 1 

Cystocloniaceae Hypnea 5 

Dasya 6 

Dasyopsis 1 

Dictyurus 1 Dasyaceae 

Heterosiphonia 2 

Delesseriaceae Nitophyllum 1 

Dumontiaceae Dudresnya 1 

Erythropeltidaceae Erythrotrichia 1 
Table A.2. (Continued). 

Appendices A-5 

Family Genus Number of Species 

Galaxaura 4 Galaxauraceae Scinaia 1 

Gelidiella 3 

Gelidiopsis 3 

Gelidium 3 Gelidiaceae 

Pterocladia 2 

Gigartinaceae Gigartina 1 

Gracilaria 16 Gracilariaceae Hydropuntia 1 

Halymeniaceae Cryptonemia 2 

Halymeniaceae Halymenia 3 

Liagoraceae Liagora 2 

Lomentariaceae Lomentaria 1 

Peyssonneliaceae Peyssonnelia 1 

Polyphysaceae Acetabularia 4 

Pyraloidea Chondria 8 

Chondrocanthus prev. Laurencia 1 

Bostrychia 2 

Botryocladia 4 

Bryocladia 1 

Bryothamnion 2 

Digenia 1 

Laurencia 8 

Polysiphonia 5 

Rhodomelaceae 

Herposiphonia 1 

Asteromenia 1 

Rhodymeniaceae Chrysymenia 2 

Rhodymenia 2 

Agardhiella 2 

Solieriaceae Neoagardhiella 0 

Solieria 2 

Ernodesmis 1 Valoniaceae Valonia 1 

Wurdemanniaceae Wurdemannia 1 

Phaeophyta 

Bibionidae Dilophus 1 

Chordariaceae Hydroclathrus 1 

Dictyopteris 2 

Dictyota 17 

Lobophora 1 

Padina 6 

Spatoglossum 1 

Dictyotaceae 

Stypopodium 1 

Ectocarpaceae Ectocarpus 1 

Pylaiellaceae Bachelotia 1 

Sargassaceae Sargassum 7 

Scarabaeoidea Hincksia 4 

Colpomenia 1 Scytosiphonaceae Rosenvingea 2 

Sphacelariaceae Sphacelaria 2 
Table A.2. (Continued). 

Appendices A-6 

Family Genus Number of Species 

Cyanophyta 

Chroococcaceae Anacystis 4 

Entophysalidaceae Entophysalis 1 

Euphorbiaceae Anabaena 1 

Hormothamnion 1 Nostocaceae Nodularia 1 

Lyngbya 0 

Oscillatoriaceae Microcoleus 3 

Oscillatoria 1 

Phormidiaceae Porphyrosiphon 1 

Rivulariaceae Calothrix 2 

Schizotrichaceae Schizothrix 2 

Total Families 57 

Total Genera 108 

Total Species 277 
Appendices A-7 

Table A.3. Genera and numbers of species of macroalgae and cyanobacteria identified within genera, by 

county, documented in the studies conducted on nearshore hardbottom of the east Florida 

coast listed in Table A.1. Genera are listed by highest frequency of occurrence among 

counties. 

Genus Brevard Indian River St. Lucie Martin Palm Beach Broward 

Number of 

Counties in which 

Species were 

Present 

Bryopsis 1 1 1 1 1 2 6 

Caulerpa 2 9 9 2 8 6 6 

Ceramium 1 4 5 3 1 5 6 

Codium 1 2 5 3 4 3 6 

Dictyota 2 4 7 2 3 12 6 

Gelidium 2 3 2 2 2 2 6 

Gracilaria 2 9 12 6 2 8 6 

Laurencia 3 4 5 2 3 4 6 

Padina 1 4 3 1 3 3 6 

Ulva 3 5 2 4 1 1 6 

Bryothamnion 2 3 2 1 3 5 

Cladaphora 1 4 2 2 2 5 

Dictyopteris 2 2 1 1 2 5 

Gelidiopsis 2 3 1 2 1 5 

Halimeda 1 2 1 4 8 5 

Hypnea 1 5 5 1 4 5 

Jania 1 2 4 1 3 5 

Sargassum 2 5 1 1 2 5 

Acanthophora 2 2 2 1 4 

Agardhiella 1 2 1 1 4 

Botryocladia 3 2 1 3 4 

Bryocladia 1 1 1 1 4 

Chaetomorpha 1 2 1 1 4 

Chondria 3 4 6 1 4 

Pterocladia 1 5 4 1 4 

Spyridia 1 2 1 1 4 

Acetabularia 1 1 3 3 

Amphiroa 2 1 3 3 

Bostrichia 1 1 1 3 

Centroceras 1 1 1 3 

Champia 1 1 1 3 

Dasya 5 2 4 3 

Erythrotrichia 1 1 1 3 

Galaxaura 1 1 1 3 

Halymenia 1 3 4 3 

Heterosiphonia 1 2 1 3 

Liagora 1 1 2 3 

Lomentaria 1 1 1 3 

Rhizoclonium 2 1 1 3 
Table A.3. (Continued). 

Appendices A-8 

Genus Brevard Indian River St. Lucie Martin Palm Beach Broward 

Number of 

Counties in which 

Species were 

Present 

Rhodymenia 2 2 1 3 

Scinaia 1 2 1 3 

Solieria 1 2 2 3 

Sphacelaria 1 1 1 3 

Udotea 2 2 6 3 

Valonia 1 1 1 3 

Acrochaetium 2 1 2 

Avrainvillea 1 2 2 

Calcareous green 1 1 2 

Calothrix 1 1 2 

Chondrocanthus 

(Laurencia) 1 1 2 

Chrysymenia 1 1 2 

Colpomenia 1 2 2 

Crouania 1 1 2 

Cryptonemia 2 1 2 

Dascycladus 1 1 2 

Eucheuma 2 3 2 

Fosliella 2 1 2 

Griffithsia 1 1 2 

Hincksia 4 2 2 

Lobophora 1 1 2 

Lyngbya 1 1 2 

Microcoleus 2 1 2 

Neomeris 1 2 2 

Nitophyllum 1 1 2 

Penicillus 1 1 2 

Rosenvingea 1 2 2 

Schizothrix 2 2 2 

Spatoglossum 1 1 2 

Spermothamnion 1 2 2 

Unidentified green 2 2 2 

Unidentified red 1 1 2 

Wrangelia 1 2 2 

Anabaena 1 1 

Anacystis 4 1 

Asparagopsis 1 1 

Asteromenia 1 1 

Bachelotia 1 1 

Batophora 1 1 

Callithamnion 2 1 

Coralline red 1 1 

Corallinaceae 1 1 
Table A.3. (Continued). 

Appendices A-9 

Genus Brevard Indian River St. Lucie Martin Palm Beach Broward 

Number of 

Counties in which 

Species were 

Present 

Cymopolia 2 1 

Dasyopsis 1 1 

Derbesia 1 1 

Dictyosphaeria 1 1 

Dictyurus 1 1 

Digenia 1 1 

Dilophus 1 1 

Dudresnya 1 1 

Ectocarpus 1 1 

Entocladia 1 1 

Entophysalis 1 1 

Ernodesmis 1 1 

Filamentous brown 1 1 

Filamentous green 1 1 

Filamentous red 1 1 

Gelidiaceae 1 1 

Gelidiella 4 1 

Gigartina 1 1 

Grateloupia 1 1 

Herposiphonia 1 1 

Hormothamnion 1 1 

Hydroclathrus 1 1 

Hydropuntia 1 1 

Lithophyllum 1 1 

Neoagardhiella 1 1 

Nodularia 1 1 

Oscillatoria 1 1 

Peyssonnelia 1 1 

Polysiphonia 1 1 

Porphyrosiphon 1 1 

Siphonocladus 1 1 

Stypopodium 1 1 

Titanophora 1 1 

Unidentified turf 1 1 

Unidentified veg. 1 1 

Ventricaria 1 1 

Wurdemannia 1 1 
Appendices A-10 Table A.4. The number of species and percent composition of sheet functional form group macroalgae in studies on nears

shading indicates similar study methods were used. High and low percent composition among similar studies a

Source Location/Tidal Zone Number of Sheet Species Total Number of Species Percent Composition Season/Year 

Continental Shelf Associates, Inc., 2005a BREVARD Inter- and subtidal combined 3 24 12.5 Summer 2005 Video tran

analysis, p

Continental Shelf Associates, Inc., 2005b PALM BEACH Shallow subtidal 2 12 16.7 2001-2004 Video tran

analysis, p

Continental Shelf Associates, Inc., 2005b PALM BEACH Subtidal artificial Video tran

analysis, p

Harris, 2006; Harris et al., 2007 MARTIN Intertidal 1 9 11.1 Summer 2001-2006 Visual tranCoastal Planning & Engineering, Inc., 2006 BROWARD Subtidal/artificial 13 100 13 Summer/Fall 2004 Transect, 

random po

Juett et al., 1976 INDIAN RIVER Intertidal 4 109 3.7 Summer 1974 Transect Phillips, 1961 ST. LUCIE Station 1, just north of St. Lucie Inlet on ocean side 6 41 14.6 SP/Fall/1957-1959 Transects Vare, 1991 PALM BEACH Intertidal 1 23 4.4 July 1987- August 1990 Transect Salmon et al., 2004 BROWARD Subtidal-preconstruction 0 6 0 2004 Visual TraHolloway-Adkins and McCarthy, 2007 BREVARD Subtidal/artificial 4 37 10.8 Summer 2007 Percent co

plates Holloway-Adkins, 2005 BREVARD Intertidal 2 24 8.3 2005 Lavage - P

Holloway-Adkins, 2001 INDIAN RIVER Intertidal 5 46 10.9 Summer 1998-2000 Lavage - PMakowski, 2004 PALM BEACH Subtidal 3 11 27.3 August-November 2003 Lavage - PWershoven and Wershoven, 1992b BROWARD Intertidal 1 10 10 YR/1992 Lavage - P

lavage Gilbert, 2005 INDIAN RIVER Subtidal 4 28 14.3 Quadrat - transects w

Cummings, 1990 PALM BEACH Intertidal artificial reef 2 10 20 Monthly Quadrat - 
Table A.4. (Continued). 

Appendices A-11 Source Location/Tidal Zone Number of Sheet Species Total Number of Species Percent Composition Season/Year 

U.S. Army Corps of Engineers, 2003a BROWARD Intertidal 2 27 7.4 AGRRA p

analysis U.S. Army Corps of Engineers, 2003a BROWARD Subtidal (species put together) (species put together) AGRRA panalysis Moffler and van Breedveld, 1979 ST. LUCIE Intertidal 6 119 5.04 September 1971- August 1974 Trawl and (includes d

Jones et al., 2004 PALM BEACH Subtidal 1 17 5.9 SP/2004 Grab samContinental Shelf Associates, Inc., 1997 ST. LUCIE Intertidal 5 29 17.2 1997 McCarthy, unpublished BREVARD 2 22 9.1 McCarthy, unpublished INDIAN RIVER Intertidal 3 42 7.1 McCarthy, unpublished ST. LUCIE Fort Pierce 8 51 15.7 McCarthy, unpublished MARTIN 1 9 11.1 McCarthy, unpublished PALM BEACH Palm Beach County 0 26 0 McCarthy, unpublished PALM BEACH North Jupiter 1 9 11.1 McCarthy, unpublished PALM BEACH South Jupiter 0 6 0 McCarthy, unpublished PALM BEACH Juno 1 8 12.5 McCarthy, unpublished PALM BEACH MacArthur Beach 0 5 0 
Appendices A-12 Table A.5. The number of species and percent composition of filamentous functional form group macroalgae in studies on 

Color shading indicates similar study methods were used. High and low percent composition among similar stu

Source Location/Tidal Zone Number of Filamentous Species Total Number of Species Percent Composition Season/Year 

Continental Shelf Associates, Inc., 2005a BREVARD Inter- and subtidal combined 5 24 20.8 Summer 2005 Video trans

analysis, pe

Continental Shelf Associates, Inc., 2005b PALM BEACH Shallow subtidal 0 12 0 2001-2004 Video trans

analysis, pe

Continental Shelf Associates, Inc., 2005b PALM BEACH Subtidal artificial Video trans

analysis, pe

Harris et al., 2007 MARTIN Intertidal 1 9 11.1 Summer 2001-2006 Visual transCoastal Planning & Engineering, Inc., 2006 BROWARD Subtidal/artificial 20 100 20 Summer/Fall 2004 Transect, B

random poi

Juett et al., 1976 INDIAN RIVER Intertidal 38 109 34.9 Summer 1974 Transect Phillips, 1961 ST. LUCIE Station 1, just north of St. Lucie Inlet on ocean side 14 41 34.2 SP/Fall/1957-1959 Transects (Vare, 1991 PALM BEACH Intertidal 2 23 8.7 July 1987- August 1990 Transect Salmon et al., 2004 BROWARD Subtidal-preconstruction 2 6 33.3 2004 Visual TranHolloway-Adkins and McCarthy, 2007 BREVARD Subtidal/artificial 13 37 35.1 Summer 2007 Percent com

plates Holloway-Adkins, 2005 BREVARD Intertidal 8 24 33.3 2005 Lavage - Pe

Holloway-Adkins, 2001 INDIAN RIVER Intertidal 16 46 34.8 Summer 1998-2000 Lavage - PeMakowski, 2004 PALM BEACH Subtidal 4 11 36.3 August-November 2003 Lavage - PeWershoven and Wershoven, 1992b BROWARD Intertidal 1 10 10 YR/1992 Lavage - Pe

necropsy Gilbert, 2005 INDIAN RIVER Subtidal 6 28 21.4 Quadrat - Ptransects w

Cummings, 1990 PALM BEACH Intertidal artificial reef 1 10 10 Monthly Quadrats - 
Table A.5. (Continued). 

Appendices A-13 Source Location/Tidal Zone Number of Filamentous Species Total Number of Species Percent Composition Season/Year 

U.S. Army Corps of Engineers, 2003a BROWARD Intertidal 7 27 25.9 AGRRA pro

analysis U.S. Army Corps of Engineers, 2003a BROWARD Subtidal (species put together) (species put together) AGRRA proanalysis Moffler and van Breedveld, 1979 ST. LUCIE Intertidal 35 119 29.4 September 1971- August 1974 Trawl and bincluded) Jones et al., 2004 PALM BEACH Subtidal 5 17 29.4 SP/2004 Grab samp

Continental Shelf Associates, Inc., 1997 ST. LUCIE Intertidal 7 51 13.7 1997 McCarthy, unpublished BREVARD 5 22 22.7 McCarthy, unpublished INDIAN RIVER Intertidal 2 42 4.8 McCarthy, unpublished ST. LUCIE Fort Pierce 7 51 13.7 McCarthy, unpublished MARTIN 5 26 19.2 McCarthy, unpublished PALM BEACH Palm Beach County 0 9 0 McCarthy, unpublished PALM BEACH North Jupiter 0 6 0 McCarthy, unpublished PALM BEACH South Jupiter 0 8 0 McCarthy, unpublished PALM BEACH Juno 0 5 0 McCarthy, unpublished PALM BEACH MacArthur Beach 0 5 0 
Appendices A-14 Table A.6. The number of species and percent composition of coarsely-branched functional form group macroalgae in stud

hardbottom. Color shading indicates similar study methods were used. High and low percent composition amoin bold. Source Location/Tidal Zone Number of Coarsely- Branched Species Total Number of Species Percent Composition Season/Year 

Continental Shelf Associates, Inc., 2005a BREVARD Inter- and subtidal combined 11 24 45.8 Summer 2005 Video tra

analysis, 

Continental Shelf Associates, Inc., 2005b PALM BEACH Shallow subtidal 8 12 66.7 2001-2004 Video tra

analysis, 

Continental Shelf Associates, Inc., 2005b PALM BEACH Subtidal artificial Video tra

analysis, 

Harris et al., 2007 MARTIN Intertidal 3 9 33.3 Summer 2001-2006 Visual traCoastal Planning & Engineering, Inc., 2006 BROWARD Subtidal/artificial 26 100 26 Summer/Fall 2004 Transect

random p

Juett et al., 1976 INDIAN RIVER Intertidal 22 109 20.2 Summer 1974 Transect Phillips, 1961 ST. LUCIE Station 1, just north of St. Lucie Inlet on ocean side 8 41 19.5 SP/Fall/1957-1959 Transects Vare, 1991 PALM BEACH Intertidal 9 23 39.1 July 1987- August 1990 Transect Salmon et al., 2004 BROWARD Subtidal-preconstruction 1 6 16.7 2004 Visual TrHolloway-Adkins and McCarthy, 2007 BREVARD Subtidal/artificial 14 37 37.8 Summer 2007 Percent c

plates Holloway-Adkins, 2005 BREVARD Intertidal 8 24 33.3 2005 Lavage - 

Holloway-Adkins, 2001 INDIAN RIVER Intertidal 12 46 26 Summer 1998-2000 Lavage - Makowski, 2004 PALM BEACH Subtidal 1 11 9.1 August-November 2003 Lavage - Wershoven and Wershoven, 1992b BROWARD Intertidal 4 10 40 YR/1992 Lavage - Gilbert, 2005 INDIAN RIVER Subtidal 9 28 32.1 Quadrat 

transects 
Table A.6. (Continued). 

Appendices A-15 Source Location/Tidal Zone Number of Coarsely- Branched Species Total Number of Species Percent Composition Season/Year 

Cummings, 1990 PALM BEACH Intertidal artificial reef 3 10 30 Monthly Quadrats U.S. Army Corps of Engineers, 2003a BROWARD Intertidal 9 27 33.3 AGRRA p

analysis U.S. Army Corps of Engineers, 2003a BROWARD Subtidal (species put together) (species put together) AGRRA panalysis Moffler and van Breedveld, 1979 ST. LUCIE Intertidal 37 119 31.1 September 1971- August 1974 Trawl and included) Jones et al., 2004 PALM BEACH Subtidal 5 17 29.4 SP/2004 Grab sam

Continental Shelf Associates, Inc., 1997 ST. LUCIE Intertidal 9 29 31.03 1997 McCarthy, unpublished BREVARD 11 22 50 McCarthy, unpublished INDIAN RIVER Intertidal 9 42 21.4 McCarthy, unpublished ST. LUCIE Fort Pierce 15 51 29.4 McCarthy, unpublished MARTIN 3 9 33.3 McCarthy, unpublished PALM BEACH Palm Beach County 8 26 30.8 McCarthy, unpublished PALM BEACH North Jupiter 2 9 22.2 McCarthy, unpublished PALM BEACH South Jupiter 2 6 33.3 McCarthy, unpublished PALM BEACH Juno 3 8 37.5 McCarthy, unpublished PALM BEACH MacArthur Beach 1 5 20 
Appendices A-16 Table A.7. The number of species and percent composition of thick-leathery functional form group macroalgae in studies o

hardbottom. Color shading indicates similar study methods were used. High and low percent composition amoin bold. Source Location/Tidal Zone Number of Thick-Leathery Species Total Number of Species Percent Composition Season/Year 

Continental Shelf Associates, Inc., 2005a BREVARD Inter- and subtidal combined 4 24 16.7 Summer 2005 Video tran

analysis, p

Continental Shelf Associates, Inc., 2005b PALM BEACH Shallow subtidal 1 12 8.3 2001-2004 Video tran

analysis, p

Continental Shelf Associates, Inc., 2005b PALM BEACH Subtidal artificial 0 12 0 Video tran

analysis, p

Harris et al., 2007 MARTIN Intertidal 4 9 44.4 Summer 2001-2006 Visual tranCoastal Planning & Engineering, Inc., 2006 BROWARD Subtidal/artificial 24 100 24 Summer/Fall 2004 Transect, 

random po

Juett et al., 1976 INDIAN RIVER Intertidal 18 109 16.5 Summer 1974 Transect Phillips, 1961 ST. LUCIE Station 1, just north of St. Lucie Inlet on ocean side 9 41 22 SP/Fall/1957-1959 Transects Vare, 1991 PALM BEACH Intertidal 4 23 17.4 July 1987- August 1990 Transect Salmon et al., 2004 BROWARD Subtidal-preconstruction 3 6 50.00 2004 Visual TraHolloway-Adkins and McCarthy, 2007 BREVARD Subtidal/artificial 5 37 13.5 Summer 2007 Percent co

plates Holloway-Adkins, 2005 BREVARD Intertidal 5 24 20.8 2005 Lavage - P

Holloway-Adkins, 2001 INDIAN RIVER Intertidal 11 46 23.9 Summer 1998-2000 Lavage - PMakowski, 2004 PALM BEACH Subtidal 2 11 18.2 August-November 2003 Lavage - PWershoven and Wershoven, 1992b BROWARD Intertidal 3 10 30.00 YR/1992 Lavage - P

necropsy Gilbert, 2005 INDIAN RIVER Subtidal 7 28 25 Quadrat - transects w
Table A.7. (Continued). 

Appendices A-17 Source Location/Tidal Zone Number of Thick-Leathery Species Total Number of Species Percent Composition Season/Year 

Cummings, 1990 PALM BEACH Intertidal artificial reef 2 10 20 Monthly Quadrats- U.S. Army Corps of Engineers, 2003a BROWARD Intertidal 3 27 11.1 AGRRA p

analysis U.S. Army Corps of Engineers, 2003a BROWARD Subtidal (species put together) (species put together) AGRRA panalysis Moffler and van Breedveld, 1979 ST. LUCIE Intertidal 30 119 25.2 September 1971- August 1974 Trawl and included) Jones et al., 2004 PALM BEACH Subtidal 4 17 23.5 SP/2004 Grab samp

Continental Shelf Associates, Inc., 1997 ST. LUCIE Intertidal 7 29 24.1 1997 McCarthy, unpublished BREVARD 4 22 18.2 McCarthy, unpublished INDIAN RIVER Intertidal 5 42 11.9 McCarthy, unpublished ST. LUCIE Fort Pierce 14 51 27.5 McCarthy, unpublished MARTIN 4 9 44.4 McCarthy, unpublished PALM BEACH Palm Beach County 7 26 26.9 McCarthy, unpublished PALM BEACH North Jupiter 1 9 11.1 McCarthy, unpublished PALM BEACH South Jupiter 0 6 0 McCarthy, unpublished PALM BEACH Juno 0 8 0 McCarthy, unpublished PALM BEACH MacArthur Beach 1 5 20 
Appendices A-18 Table A.8. The number of species and percent composition of jointed-calcareous functional form group macroalgae in stu

hardbottom. Color shading indicates similar study methods were used. High and low percent composition amin bold. Source Location/Tidal Zone Number of Jointed- Calcareous Species Total Number of Species Percent Composition Season/Year 

Continental Shelf Associates, Inc., 2005a BREVARD Inter- and subtidal combined 0 24 0 Summer 2005 Video tra

analysis, 

Continental Shelf Associates, Inc., 2005b PALM BEACH Shallow subtidal 1 12 8.3 2001-2004 Video tra

analysis, 

Continental Shelf Associates, Inc., 2005b PALM BEACH Subtidal artificial 0 12 0 Video tra

analysis, 

Harris et al., 2007 MARTIN Intertidal 0 9 0 Summer 2001-2006 Visual traCoastal Planning & Engineering, Inc., 2006 BROWARD Subtidal/artificial 17 100 17 Summer/Fall 2004 Transect

random p

Juett et al., 1976 INDIAN RIVER Intertidal 7 109 6.4 Summer1974 Transect Phillips, 1961 ST. LUCIE Station 1, just north of St. Lucie Inlet on ocean side 2 41 4.9 SP/Fall/1957-1959 TransectVare, 1991 PALM BEACH Intertidal 5 23 21.7 July 1987- August 1990 Transect Salmon et al., 2004 BROWARD Subtidal-preconstruction 0 6 0 2004 Visual TrHolloway-Adkins and McCarthy, 2007 BREVARD Subtidal/artificial 1 37 2.7 Summer 2007 Percent c

plates Holloway-Adkins, 2005 BREVARD Intertidal 1 24 4.2 2005 Lavage - 

Holloway-Adkins, 2001 INDIAN RIVER Intertidal 2 46 4.4 Summer 1998-2000 Lavage - Makowski, 2004 PALM BEACH Subtidal 1 11 9.1 August-November 2003 Lavage - Wershoven and Wershoven, 1992b BROWARD Intertidal 1 10 10 YR/1992 Lavage - 

necropsy Gilbert, 2005 INDIAN RIVER Subtidal 2 28 7.1 Quadrat transects 
Table A.8. (Continued). 

Appendices A-19 Source Location/Tidal Zone Number of Jointed- Calcareous Species Total Number of Species Percent Composition Season/Year 

Cummings, 1990 PALM BEACH Intertidal artificial reef 2 10 20 monthly Quadrats U.S. Army Corps of Engineers, 2003a BROWARD Intertidal 5 27 18.5 AGRRA 

analysis U.S. Army Corps of Engineers, 2003a BROWARD Subtidal (species put together) (species put together) AGRRA analysis Moffler and van Breedveld, 1979 ST. LUCIE Intertidal 7 119 5.9 September 1971- August 1974 Trawl anincluded) Jones et al., 2004 PALM BEACH Subtidal 0 17 0 SP/2004 Grab sam

Continental Shelf Associates, Inc., 1997 ST. LUCIE Intertidal 2 29 6.7 1997 McCarthy, unpublished BREVARD 0 22 0 McCarthy, unpublished INDIAN RIVER Intertidal 1 42 2.4 McCarthy, unpublished ST. LUCIE Fort Pierce 4 51 7.8 McCarthy, unpublished MARTIN 1 9 11.1 McCarthy, unpublished PALM BEACH Palm Beach County 6 26 23.1 McCarthy, unpublished PALM BEACH North Jupiter 2 9 22.22 McCarthy, unpublished PALM BEACH South Jupiter 1 6 16.7 McCarthy, unpublished PALM BEACH Juno 1 8 12.5 McCarthy, unpublished PALM BEACH MacArthur Beach 1 5 20 
Appendices A-20 Table A.9. The number of species and percent composition of crustose functional form group macroalgae in studies on n

Color shading indicates similar study methods were used. High and low percent composition among similar st

Source Location/Tidal Zone Number of Crustose Species Total Number of Species Percent Composition Season/Year 

Continental Shelf Associates, Inc., 2005a BREVARD Inter- and subtidal combined 0 24 0 Summer 2005 Video tra

analysis, 

Continental Shelf Associates, Inc., 2005b PALM BEACH Shallow subtidal 0 12 0 2001-2004 Video tra

analysis, 

Continental Shelf Associates, Inc., 2005b PALM BEACH Subtidal artificial 0 12 0 Video tra

analysis, 

Harris et al., 2007 MARTIN Intertidal 0 9 0 Summer 2001-2006 Visual traCoastal Planning & Engineering, Inc., 2006 BROWARD Subtidal/artificial 0 100 0 Summer/Fall 2004 Transect

random p

Juett et al., 1976 INDIAN RIVER Intertidal 2 109 1.8 Summer 1974 Transect Phillips, 1961 ST. LUCIE Station 1, just north of St. Lucie Inlet on ocean side 0 41 0 SP/Fall/1957-1959 TransectVare, 1991 PALM BEACH Intertidal 0 23 0 July 1987- August 1990 Transect Salmon et al., 2004 BROWARD Subtidal-preconstruction 0 6 0 2004 Visual TrHolloway-Adkins and McCarthy, 2007 BREVARD Subtidal/artificial 0 37 0 Summer 2007 Percent c

plates Holloway-Adkins, 2005 BREVARD Intertidal 0 24 0 2005 Lavage - 

Holloway-Adkins, 2001 INDIAN RIVER Intertidal 0 46 0 Summer 1998-2000 Lavage - Makowski, 2004 PALM BEACH Subtidal 0 11 0 August-November 2003 Lavage - Wershoven and Wershoven, 1992b BROWARD Intertidal 0 10 0 YR/1992 Lavage - 

necropsy Gilbert, 2005 INDIAN RIVER Subtidal 0 28 0 Quadrat transects 

Cummings, 1990 PALM BEACH Intertidal artificial reef 0 10 0 Monthly Quadrats 
Table A.9. (Continued). 

Appendices A-21 Source Location/Tidal Zone Number of Crustose Species Total Number of Species Percent Composition Season/Year 

U.S. Army Corps of Engineers, 2003a BROWARD Intertidal 5 27 18.5 AGRRA 

analysis U.S. Army Corps of Engineers, 2003a BROWARD Subtidal (species put together) (species put together) AGRRA analysis Moffler and van Breedveld, 1979 ST. LUCIE Intertidal 0 119 0 September 1971- August 1974 Trawl anincluded) Jones et al., 2004 PALM BEACH Subtidal 0 17 0 SP/2004 Grab sam

Continental Shelf Associates, Inc., 1997 ST. LUCIE Intertidal barrier island that separates Indian River Lagoon from Atlantic 0 29 0 1997 McCarthy, unpublished BREVARD 0 22 0 McCarthy, unpublished INDIAN RIVER Intertidal 0 42 0 McCarthy, unpublished ST. LUCIE Fort Pierce 0 51 0 McCarthy, unpublished MARTIN 0 9 0 McCarthy, unpublished PALM BEACH Palm Beach County 0 26 0 McCarthy, unpublished PALM BEACH North Jupiter 0 9 0 McCarthy, unpublished PALM BEACH South Jupiter 0 6 0 McCarthy, unpublished PALM BEACH Juno 0 8 0 McCarthy, unpublished PALM BEACH MacArthur Beach 0 5 0 
Appendices B-1 

Appendix B
Appendices B-2 

Table B.1. Presence/absence of invertebrates encountered in available studies of hardbottom habitats in 

six counties along the east Florida coast. ● denotes species identified in at least one study 

within noted county; blank cell denotes species not encountered in any study in noted county. 

Note that species are listed as identified in the respective study, and that those highlighted in 

yellow were not used to determine species counts for the taxonomic groups listed in 

Chapter 4. 

Taxon Brevard Indian River St. Lucie Martin Palm Beach Broward 

Polychaetes 

Anaitidae longipes ● 

Anamobaea oerstedii ● ● 

Arabella sp. ● 

Aricidea sp. ● 

Armandia agilis ● 

Axiothella sp. ● 

Brachioma nigromaculata ● ● ● 

Clymenella torquata ● 

Decamastus sp. ● 

Dispio unicata ● 

Doruilleidae sp. ● 

Euclymene sp. ● 

Eupolymnia crassicornis ● ● 

Exogone atlantica ● 

Genetyllis sp. ● 

Glycera sphyrabrancha ●

Goniadella sp. ● 

Haplosyllis spongicola ● 

Hermodice carunculata ●

Kefersteinia cirrata ● ● 

Leitoscoplos fragilis ● ● 

Lumbrneris sp. ● 

Macroclymene sp. ● 

Magelona sp. ● 

Marphysa sanquinea ●

Mediomastus californiensis ●

Megelonna sp. ● 

Myriochele oculata ●

Naineris grubei ●

Nematonereis hebes ●

Nepthys squamosa ●

Nereis falsa ●

Nereis lamellose ●

Nereis pelagica ●

Notomastus sp. ● 

Odontosyllis enopla ●

Pectinaria gouldii ●

Peresiella sp. ● 

Pherusa inflata ●

Phragmatopoma lapidosa (caudata) ● ● ● ● ● ●

Pista sp. ● 

Prionospio cirrifora ● 
Table B.1. (Continued). 

Appendices B-3 

Taxon Brevard Indian River St. Lucie Martin Palm Beach Broward 

Prionospio cristata ● 

Prionospio fallax ●

Prionospio pygmaea ●

Pseudovermilia occidentalis ●

Sabella sp. A ● 

Sabella sp. B ● 

Sabellastarte magnifica ● 

Sabellidae ● 

Saccocirrus sp. ● 

Salmacina sp. ● 

Scolepis squamata ● 

Sabellaridae Species A ● 

Serpulidae Species A ● 

Serpulidae Species B ● 

Sicyonia brevirostris ● 

Spiophanes bombyx ●

Spiophanes missionensis ●

Spirobranchus giganteus ● 

Terribellidae sp. ● 

Unknown species ● 

Websterinereis tridentate ● 

Anthozoans 

Acropora cervicornis ●

Acropora palmata ●

Agaricia agaricites ●

Agaricia fragilis ●

Agaricia sp. ● 

Cladocora arbuscula ● ●

Colpophyllia natans ● ●

Dendrogyra cylindrus ●

Dichocoenia stokesii ● ● ●

Diploria clivosa ● ● ●

Diploria labyrinthiformis ●

Diploria sp. ● 

Diploria strigosa ● ● ● ●

Favia fragum ● ● ●

Isophyllia sinuosa ● ● ●

Isophyllia sp. ● 

Madracis decactis ● ● ●

Meandrina meandrites ● ● ●

Montastraea annularis ●

Montastraea franksi ●

Montastraea cavernosa ● ● ●

Montastraea faveolata ● ● ●

Mycetophyllia aliciae ● ● ●

Mycetophyllia danaana ●

Oculina diffusa ● ● ● ● ●

Oculina robusta ● ●
Table B.1. (Continued). 

Appendices B-4 

Taxon Brevard Indian River St. Lucie Martin Palm Beach Broward 

Oculina varicosa ● ● ● ● ●

Phyllangia americana ● ● ● ● ●

Porites astreoides ● ● ●

Porites porites ● ● ●

Porites sp. ● 

Scolymia sp. ● 

Siderastrea radians ● ● ● ● ●

Siderastrea siderea ● ● ● ● ●

Siderastrea sp. ● ● ● 

Solenastrea bournoni ● ● ● ●

Solenastrea hyades ● ● ●

Stephanocoenia intersepta ● ● ●

Bunodosoma caveranta ● ● ●

Actinia bermudnesis ● ● ●

Diadumene leucolena ● ● ●

Palythoa caribaeorum ● ● ●

Zoanthus sp. ● ●

Zoanthus pulchellus ● ●

Briareum asbestinum ● ● ●

Carijoa riisei ● ● ● ●

Erythropodium caribaeorum ● ●

Eunicea calyculata ● ●

Eunicea sp. ● ● ●

Gorgonia sp. ● 

Gorgonia ventalina ● ● ●

Leptogorgia hebes ● ● ● ● ●

Leptogorgia virgulata ● ● ● ● ●

Muricea sp. ● ● ●

Plexaura flexuosa ● ●

Plexaurella grisea ● ● ●

Plexaurella sp. ● 

Pseudopterogorgia americana ● ●

Pseudopterogorgia sp. ● ● ● 

Pterogorgia anceps ● ● ●

Pterogorgia citrina ● ● ●

Pterogorgia guadalupensis ● ● ●

Pterogorgia sp. ● ● 

Unidentified sea rod ● 

Zoanthus sp. ● 

Sponges

Agelas conifer ● 

Agelas schmidti ● 

Agelas sp. ● 

Aiolochroia crassi ● 

Amphimedon compressa ● ● 

Anthosigmella varians ● ● ●

Aplysina cauliformis ● 

Aplysina fistularis ● ● 
Table B.1. (Continued). 

Appendices B-5 

Taxon Brevard Indian River St. Lucie Martin Palm Beach Broward 

Aplysina sp ● 

c.f. Dysidea etheria ●

Calcispongidae sp. ● 

Callyspongia sp. ● ● 

Callyspongia vaginalis ● 

Chondrilla nucula ● 

Cinachyra alloclado ● ● 

Cinachyra sp. ● ● 

Cliona celata ● 

Cliona delitrix ● ● 

Cliona lampa ● ● ● ● 

Cliona lampa forma flavida ● 

Cliona lampa forma lampa ● 

Cliona sp. ● ● ● ● 

Cliona viridis ● 

Desmapsamma sp. ● 

Diplastrella sp. ● 

Encrusting sponges ● ● 

Eurospongia rosea ● 

Euryspongia rasea ● 

Geodia sp. ● 

Haliclona sp. ● 

Halicona(purple) ● 

Halicona(red) ● 

Haloclona rubens ● 

Holopsamma sp. ● 

Iotrochota birotulata ● 

Ircinia campana ● ● ● 

Ircinia fasciculata ● 

Ircinia felix ● 

Ircinia strobilina ● ● 

Ircinia variabilis ● 

Lotrochota birotulata ● 

Microciona sp. ● ● 

Microciona spinosa ● 

Monanchora unguifera ● 

Niphates digitalis ● 

Niphates erecta ● ● 

Niphates sp. ● ● 

Niphates(purple) ● ●

Plakortis angulospiculatus ●

Poecilosclerida spp. ● 

Pseudoaxinella lunaecharta ● ● ● 

Sigmadocia caerulea ● 

Spheciospongia vesparium ● ● 

Spheciospongia vesparsa ● 

Spinosella plicifera ● 

Spinosella vaginalis ● 
Table B.1. (Continued). 

Appendices B-6 

Taxon Brevard Indian River St. Lucie Martin Palm Beach Broward 

Spirastrella coccinea ● 

Spongia sp. ● 

Teichaxinella sp. ● 

Tethya sp. ● ● 

Ulosa reutzleri ● 

Unidentified orange sponge ● 

Unidentified red sponge ● ● 

Unidentified sponge1 ● ● ● 

Unidentified tube-type ● 

Unidentified yellow sponge ● ● 

Verongula sp. ● 

Xestospongia muta ● 

Hydrozoans 

Aglaophenia latecarinata ● 

Aglaophenia sp. ● 

Campanularia marginata ● 

Campanularia sp. ● 

Eudendrium ramosum ● 

Eudendrium sp. ● 

Feather hydroids ●

Hydroida undet. sp. ● 

Lytocarpus philippinus ● 

Millepora alcocornis ● ● ●

Millepora complanata ● 

Millepora sp. ● 

Obelina hyanlina ● 

Pennaria sp. ● 

Sertularia flowersi ● 

Sertuarella sp. ● 

Sertularella speciosa ● 

Thryroscuphus ramosus ● 

Thyroscyphus marginatus ● 

Thyryoscyphus sp. ● 

Unident. branching hydroid ● 

Unidentified hydroid ● ● 

Unidentified star-shaped ● 

Yellow branch hydroid ● 

Platyhelminthes 

Unid Turbellaria sp. ● 

Polycladia sp.??? ● 

Nematoda 

Unidentified sp. ● 

Sipuncula 

Unidentified sp. ● 

Unidentified sp. 1 ●

Unidentified sp. 2 ●

Unidentified sp. 3 ●

Bryozoans 
Table B.1. (Continued). 

Appendices B-7 

Taxon Brevard Indian River St. Lucie Martin Palm Beach Broward 

Aetea sica ● ● 

Aetea truncate ● ●

Aeverrillia armata ● ●

Aimulosia pusilla ● 

Aimulosia uvulifera ●

Alcyondium polypylum ●

Alcyonidium capronae ●

Alderina smitti ●

Amathia alternata ● ● 

Amathia distans ● 

Amathia vidovici ● ● 

Anguinella palmata ● ●

Antropora leucocypha ● ●

Bartensia minuata ● 

Beania hirtissima ●

Beania intermedia ●

Beania klugei ● ● 

Beania mirabilis ● 

Bellulophora bellula ●

Bowerbankia gracilis ● ● 

Bowerbankia imbricata ● 

Bowerbankia maxima ● ● 

Bryozoan sp. ● 

Bugula minima ●

Bugula neritina ● ● 

Bugula stolonifera ● ● ● 

Bugula turrita ● 

Bugula uniserialis ● ● 

Caulibugula pearsei ● 

Celleporella carolinensis ●

Celleporina hassalli ●

Crisia elongata ● ● 

Cryptosula pallasiana ● ●

Cupuladria doma ● 

Cymulopora uniserialis ●

Discoporella umbellata depressa ●

Discoporella umbellate ●

Disporella plumose ●

Drepanophora torquata ●

Electra bellula ●

Escharina pesanseris ●

Exechonella antillea ● ● 

Floridina parvicella ● 

Floridinella typical ● ● 

Hippaliosina rostrigera ● 

Hippoporina verrilli ●

Hippothoa balanophila ●

Membranipora arborescens ●
Table B.1. (Continued). 

Appendices B-8 

Taxon Brevard Indian River St. Lucie Martin Palm Beach Broward 

Membranipora savarii ● ● 

Membranipora triangularis ● 

Membranipora tuberculata ●

Microporella umbracula ●

Nolella stipata ● ● 

Parasmitina nitida ● 

Parasmitina signata ●

Pasythea tulipifera ●

Phylactella ais ●

Phylactellipora aviculifera ●

Reginella repangulata ●

Retevirgula caribbea ●

Savignyella lafontii ● ● 

Schizoporella cornuta ● ●

Schizoporella rugosa ● 

Schizoporella unicornis ●

Scrupocellaria regularis ●

Spathipora brevicauda ●

Stylopoma spongites ●

Sundanella sibogae ●

Synnatum aegyptiacum ● ● 

Thalamoporella floridana ● ●

Trematooecia psammophila ● 

Trypostega venusta ●

Tubulanus riceae ●

Valkeria atlantica ●

Vibracellina laxibasis ●

Vittaticella contei ● ● 

Vittaticella uberrima ● 

Watersipora sp. ● ● 

Watersipora subovoidea ● ● ● 

Zoobotryon verticillatum ● ● 

Barnacles 

Tetraclita s. stalactifera ● ● ● ● ● ●

Balanus t. antillensis ● ● 

Megabalanus cocopoma ● ● ● 

Balanus eburneus ● ● ● ● ● ●

Balanus amphitrite ● ● ● ● ● ●

Chthamalus fragilis ● ● ● ● ● ●

Balanus improvisus ● ●

Chthamalus stellatus ● ●

Balanus venustus ● 

Tunicates 

Phallusia nigra ● ● ●

Mogula sp. ● 

Botryllus planus ● ● ● ●

Botrylloides nigrum ● ● ● ●

Didemnum candidum ● 
Table B.1. (Continued). 

Appendices B-9 

Taxon Brevard Indian River St. Lucie Martin Palm Beach Broward 

Didemnum sp. ● 

Eudistoma capsilatum ● 

Eudistoma obscuratum 

Diplosoma macdonaldi ● 

Diplosoma sp. ● 

Distaplia bermudensis ● 

Aplidium lobatum(?) ●

Perophora viridis ●

Trididemnum orbiculatum ●

Trididemnum savignii ●

Ecteinascidea turbinata ●

Didemnum antillensis ● ● ● ● ● ●

Echinoderms 

Echinometra lucunter ● ● ● ● ● ●

Arbacia punctulata ● ● ● 

Eucidaris tribuloides ● ● ● ● ●

Diadema antillarum ● ● ● ●

Lytechinus variagatus ● ● ● ●

Tripneustes ventriculosus ● ● ● ● ●

Holothuria grisea ● ● ● ● 

Davidaster rubiginosa ● 

Nemaster grandis ● 

Amphioholis squamata ● 

Ophiuroidea sp. ● 

Ophiothrix suensonii ● ● 

Dendrochirotida sp. ● 

Cucumariidae sp. ● 

Isostichopus badyonotus ● 

Isostichopus sp. ● 

Unidentified sp. ● 

Nemerteans 

Amphiporus ochraceous ● 

Amphiporus texanus ● 

Baseodiscus delineatus ● ● ●

Emplectonema osceolai ●

Nemertopsis bivittata ● ● ●

Prosorhochmus americanus ● ●

Prosorhochmus belizeanus ● 

Tetrastemma enteroplecta ● ● 

Tetrastemma merula ● ● 

Tetrastemma worki ● ● 

Tubulanus pellucidus ● 

Zygonemertes cocacola ● 

Zygonemertes simonae ●

Zygonemertes virescens ●

Crustaceans 

Metapenaeopsis goodei ● 

M. smithi ● 
Table B.1. (Continued). 

Appendices B-10 

Taxon Brevard Indian River St. Lucie Martin Palm Beach Broward 

Penaeus sp. ● 

Sicyonia dorsalis ● 

Neopontonides beaufortensis ● 

Periclimenes americanus ● 

Alpheus armillatus ● 

A. bouvieri ● ● 

A. formosus ● 

A. heterochaelis ● 

A. malleator ● 

A. normanni ●

A. nuttingi ●

A. paracrinitl/s ●

A. peasei ●

A. thomasi ● 

A. viridari ● 

Metalpheus rostratipes ●

Synalpheus brevicarpus ● 

S. cf. cl/racaoensis ● 

S. fritzmuelleri ● ●

S. minl/s ● 

S. townsendi ●

Synalpheus sp. A ● 

Lysmata intermedia ●

L. wurdemanni ●

Processa fimbriata ●

Microprosthema semilaeve ● 

Panulirus argus ● 

Upogebia aftinis ● 

Megalobrachium poeyi ●

M. soriatum ● 

Pachycheles monilifer ● ● ●

Petrolisthes armatus ● 

P. galathinus ● ● 

Calcinus tibicen ●

Clibanarius antillensis ●

C. sclopetarius ●

C. tricolor ●

C. vittatus ● 

Paguristes tortugae ● 

Petrochirus diogenes ● 

Pagurus carolinensis ● 

P. pollicaris ●

Lepidopa benedicti ● 

Emerita talpoida ● 

Uhlias limbatus ● 

Arenaeus cribrarius ● 

Callinectes ornatus ● 

C. similis ● 
Table B.1. (Continued). 

Appendices B-11 

Taxon Brevard Indian River St. Lucie Martin Palm Beach Broward 

Cronius ruber ● 

Portunus gibbesii ●

Platyactaea setigera ●

Cataleptodius floridanus ● 

Eriphia gonagra ● 

Eurypanopeus abbreviatus ● ● 

E. depressus ● 

E. dissimilis ● 

Hexapanopeus angustifrons ● 

H. paulensis Rathbun ● 

Menippe mercenaria ● ● 

M. nodifrons ● ● ● 

Micropanope granulimanus ● 

Panopeus bermudensis ● ● 

P. herbstii ● 

P. occidentalis ● 

Pilumnus dasypodus ● ●

P. lacteus Stimpson ● 

P. sayi Rathbun ●

Xantho denticulatus ● 

Pachygrapsus transverses ● ● ● 

Plagusia depressa ● ● 

Gecarcinus lateralis ●

Ocypode quadrata ● 

Acanthonyx petiverii ● 

Chorinus heros ●

Epialtus bituberculatus ●

E. dilatatus ●

Macrocoeloma subparallelum ●

Microphrys bicornutus ●

M. antillensis ● 

Mithrax acuticornis ● 

M. coryphe ● 

M. forceps ● ● 

M. hispidus ● 

M. pleuracanthus ●

M. ruber ● 

M. verrucosus ● 

Pelia mutica ●

Pitho lherminieri ●

Podochela riisei ● 

P. sidneyi ●

Stenorhynchus seticornis ● 

Gonodactylus bredini ● 

G. oerstedii ●

G. spinulosus ●

Pagurus brevidactylus ●

Paguristes hummi ● 
Table B.1. (Continued). 

Appendices B-12 

Taxon Brevard Indian River St. Lucie Martin Palm Beach Broward 

Tanystylum orbiculare ● 

Periclimenes americanus ● 

Paradella dianae ● ● 

Sphaeroma walkeri ● ● 

Paranthura infundibulata ● ● 

Jaeropsis sp. ● ● 

Excorallana spp. ● ● 

Hyale perieri ● ● 

Elasmopus pectinicrus ● ● 

Ampithoe pollex ● ● 

Corophium acutum ● ● 

Ericthonius sp. ● ● 

Jassa cf. falcata ● ● 

Ampithoe longimana ● ● 

Stenothoe georgiana ● ● 

Podocerus brasiliensis ● ● 

Cymadusa compta ● ● ● 

Percnon gibbesi ● 

Pagurus carolinensis ● 

Mollusca 

Aplysia dactylomela ● ● 

Anachis avara ● 

Anachis floridana ● ● ● 

Anachis lafresnayi ● ● 

Arca zebra ● 

Astrea tuber ● ● ● 

Atrina ridgida ● 

Barleeia tincta ● ● 

Cerithium litteratum ● 

Chama macreroohvilla ● 

Chlamus senits) ● 

Costoanachis sp. ● 

Lima pellucid ● 

Loligo pealeii ? ● 

Cyprea sp. ● ● ● 

Erato maugeriae ● ● 

Fissurella barbadensis ● ● 

Lamellaria perspicua ● ● 

Lucapinella limatula ● ● 

Mitrella argus ● ● 

Morula didyma ● 

Nudibranchia sp. ● ● 

Octopus vulgaris ● 

Petaloconchus sp. ● 

Petaloconchus varians ● ● ● 

Phidiana lynceus ● ● 

Pisania tinctus ● ● 

Stramonita (Thais) haemostoma ● ● 
Table B.1. (Continued). 

Appendices B-13 

Taxon Brevard Indian River St. Lucie Martin Palm Beach Broward 

floridana 

Thais rustica ● ● 

Tricolia affinis pterocladica ● ● 

Aplysia dactylomela ● ● 

Littorina sp. ● 

Iselica fenestrate ● ● 

Barbatia dominensis ● ● 

Isognomon radiatus ● ● 

Musculus lateralis ● ● 

Ostrea equestris ● ● ● 

Ostrea sp. ● 

Sphenia antillensis ● ● 

Anomia sp. ● ● 

Pteria sp. ● ● 

Pteria columbus ● 

Hiatella solida ● 

Octopus briareus ● 

Octopus joubini ● 

Sepioteuthis sepioidea ● ● 

Phoronida

Unidentified species ● ● 

Studies reviewed to compile this species list were Nelson (1989), Coastal Science Associates, Inc. (2000), M. Rice, 

personal communication, Rudolph (1977), Gore et al. (1978), Nelson (1988), Nelson and Demetriades (1992), 

Watanabe (2002), Watanabe and Young (2006), McCarthy (2001). Eckelbarger (1976), Reed et al. (1982), Reed 

(1982), Reed and Mikkelsen (1987), SFCREMP (2007), Prekel at al. (2008), Jaap (1984), Vare (1991), J. Beal, pers. 

comm., McCarthy, pers. obs., Coastal Planning & Engineering, Inc. (2006), Brooke and Young (2005), Vargas-Angel 

et al. (2006), Brooke (2002), Cummings (1994), McCarthy (2006, 2008), Walton Smith et al. (1950), Multer and 

Milliman (1967), Van Montfrans (1981), Applied Biology, Inc. (1979), Petuch (1987), Coastal Science Associates, Inc. 

(2000), Continental Shelf Associates, Inc. (1984), Continental Shelf Associates, Inc. (2002a,b, 2003, 2004, 2005b, 

2006), Department of Environmental Resources Management (1993, 1994), Harris and Lee (2006), U.S. Army Corps 

of Engineers (2003b), U.S. Army Corps of Engineers (2002), Kirtley (1966), Moyer et al. (2003), and Winston (1982). 
Appendices B-14 

Table B.2. Molluscan community composition and total number of individuals per species collected from 

41 quantitative samples of Oculina varicosa coral on four reefs off the east Florida coast. 

Species characterizations (MOE, FOOD, ZOOG, Z) were taken from the literature (sources 

listed in footnotes) and do not reflect the results of this study unless otherwise noted. 

ZOOG and Z were assigned only to fully identified species. (― = data unavailable or 

unknown; * or † preceding species name = abundant [>100 N], or common [10-100 N] 

species, respectively; * or - in station columns = presence or absence, respectively, in 

qualitative samples.) (From: Reed and Mikkelsen, 1987). 

MOE* FOOD† ZOOG‡ Z§ 6 m 27 m 42 m 80 m Total 

Class Gastropoda 

 Subclass Prosobranchia 

 Order Archaeogastropoda 

Scissurellidae 

 Scissurella proxima (Dall, 1927) F D TC A/B/C 1 0 0 0 1 

 Fissurellidae 

 Diodora arcuata (Sowerby, 1862) F H T A 0 0 0 4 4 

 †D. cf. arcuata (Sowerby, 1862) F H --- --- 0 0 II 16 27 

 D. cayenensis (Lamarck, 1822) F H TN2 A 3 0 0 0 3 

 D. cf. cayenensis (Lamarck, 1822) F H --- --- 0 0 1 0 1 

 D. dysoni (Reeve, 1850) F H T B 1 0 0 0 1 

 D. cf. dysoni (Reeve, 1850) F H --- --- 3 0 0 0 3 

 D. cf. /isteri (Orbigny, 1842) F H --- --- 0 0 4 0 4 

 D. sp. F H --- --- 7 0 0 1 8 

 Lucapinella /imatula (Reeve, 1850) F H TC A/B 1 0 0 0 I 

 Puncturella sp. F H --- --- --- --- * --- * 

 Diodorinae sp. I F H --- --- 0 1 1 5 7 

 Diodorinae sp. 2 F H --- --- 0 0 3 4 7 

 Diodorinae sp. 3 F H --- --- 0 0 0 5 5 

 Diodorinae sp. 4 F H --- --- 1 0 0 0 1 

 Diodorinae sp. 5 F H --- --- 0 0 1 3 4 

Trochidae 

 Calliostoma euglyptum (A. Adams, 1854)║ F H/O TC A 0 0 8 0 8 

 C. jujubinum (Gmelin, 1791)║ F D/O TC A/B --- --- * --- * 

 C. pulchrum (C.B. Adams, 1850)║ F D/O TC A/B/C 0 0 1 4 5 

 †C. roseolum (Dall, 1880)║ F D/O TC A/B/C 0 0 0 15 15 

 †C. sp. F --- --- --- 1 0 23 48 72 

Clyclostrematidae 

 †Arene bairdii (Dall, 1889) F H TC B/C 0 0 0 37 37 

 A. sp. F H --- --- 0 0 0 2 2 

Turbinidae 

Turbo castanea (Gmelin, 1791) F H TC A 0 0 1 0 1 

Phasianellidae 

Tricolia affinis pterocladica (Robertson, 1958) F D/H T A 1 0 0 0 1 

T. thalassicola (Robertson, 1958) F D/H TC A/B 0 0 5 0 5 

T. sp. F D/H --- --- 1 0 3 0 4 

Order Mesogastropoda 

Truncatellidae 

Truncatella cf. scalaris (Michaud, 1830) F ?D/H --- --- 1 0 0 0 1 

Rissoidae 

Alvania auberiana (Orbigny, 1842) F D/H TC A 0 0 4 3 7 

Barleeia sp. F D/H --- --- 1 0 0 0 1 

* Rissoina bouryi (Desjardin, 1949) F H T --- 102 0 0 0 102 

Caecidae 

 †Caecurn pu/chellurn (Stimpson, 1851) F H TN4 A 11 0 0 0 11 

 C. sp. F H --- --- 3 0 0 0 3 

Meioceras cubitatum (Folin, 1868) F H TC A/B 0 0 3 0 3 

M. nitidurn (Stimpson, 1851) F H T A 2 0 0 0 2 

Vitrinellidae 

Macrornphalina palmalitoris (Pilsbry and McGinty, 

1950) F --- T B --- --- * --- * 

 *Parviturboides interruptus (C.B. Adams, 1850) F D/H TC A/B 1,103 0 0 1 1,104 

Cerithiopsidae 

Cerithiopsis bicolor (C.B. Adams, 1845) ?S---S ?P T --- 0 0 2 0 2 

C. flavurn (C.B. Adams, 1850) ?S---S ?P T --- 0 0 0 7 7 

C. io (Dall and Bartsch, 1911) ?S---S ?P T --- 5 0 0 0 5 

C. sp. 3 ?S---S ?P --- --- 0 0 0 9 9 

C. sp. 4 ?S---S ?P --- --- 0 0 1 78 79 

C. sp. 5 ?S---S ?P --- --- 0 0 1 51 52 
Table B.2. (Continued). 

Appendices B-15 

MOE* FOOD† ZOOG‡ Z§ 6 m 27 m 42 m 80 m Total 

C. sp. 6 ?S---S ?P --- --- 0 0 2 4 6 

C. sp. 7 ?S---S ?P --- --- 1 0 0 0 1 

C. sp. 8 ?S---S ?P --- --- 1 0 0 0 1 

C. sp. 9 ?S---S ?P --- --- 0 0 2 0 2 

C. Sp. 10 ?S---S ?P --- --- 0 0 0 2 2 

C. sp. ?S---S ?P --- --- 0 0 0 1 1 

Seila adamsi (H.C. Lea, 1845) ?S---S ?P TN3 A/B 0 0 6 2 8 

 †S. sp. 1 ?S---S ?P --- --- 6 0 6 6 18 

 S. sp. 2 ?S---S ?P --- --- 6 0 2 1 9 

Turritellidae 

 †Vermicularia spirata (Philippi, 1836) E F TN3 A 0 0 18 71 89 

 Turritellidae sp. E F --- --- 0 0 1 0 1 

Vermetidae 

Serpulorbis decussatus (Gmelin, 1791) .E F TC A 0 0 5 1 6 

 Vermetidae sp. E F --- --- 0 0 2 0 2 

Architectonicidae 

 Architectonica peracuta (Dall, 1889) S---A C TC B 0 0 0 3 3 

 †Heliacus bisu/catus (Orbigny, 1842) S---A P/?O TC A/B/C 0 0 0 10 10 

Mathildidae 

Mathilda barbadensis (Dall, 1889) F ?C T B 0 0 0 1 1 

Triphoridae 

*Metaxia rugulosa (C.B. Adams, 1850) S---S P T A/B 1 0 0 277 278 

M. taeniolata (Dall, 1889) ?S---S ?P M A/B --- --- * * * 

Triphora cf. lilacina (Dall, 1889) ?S---S ?P --- --- 0 0 0 1 1 

T. cf. nigrocincta (C.B. Adams, 1839) ?S---S ?P --- --- 1 0 0 0 1 

 T. sp. I ?S---S ?P --- --- 1 0 7 0 8 

 †T. sp. 2 ?S---S ?P --- --- 0 0 5 61 66 

 T: sp. 3 ?S---S ?P --- --- 0 0 1 1 2 

 T. sp. 4 ?S---S ?P --- --- 1 0 0 0 1 

 T. sp. 5 ?S---S ?P --- --- 0 0 1 8 9 

 T. sp. 6 ?S---S ?P --- --- 0 0 0 3 3 

 T. sp. 7 ?S---S ?P --- --- 0 0 0 2 2 

 T. sp. 8 ?S---S ?P --- --- 0 0 0 1 1 

 T. sp. 9 ?S---S ?P --- --- 0 0 0 2 2 

 †T. sp. ?S---S ?P --- --- 0 0 1 0 10 

Pyramidellidae 

 † Fargoa bushiana (Bartsch, 1909) ?S---M, P ?P M3 A 15 0 8 36 59 

F. dianthophila (Wells and Wells, 1961) S---P P M3 A 2 0 5 1 8 

 †Odostomia# somersi (Verrill and Bush, 1900) ?S---M, P ?P T --- 0 0 0 18 18 

 O. sp. 2 ?S---M, P ?P --- --- 2 0 1 1 4 

 O. sp. 3 ?S---M, P ?P --- --- 2 0 1 2 5 

 †O. sp. 4 ?S---M, P ?P --- --- 0 0 4 10 14 

 †O. sp. 5 ?S---M, P ?P --- --- 0 0 1 67 68 

 †O. sp. 6 ?S---M, P ?P --- --- 22 0 0 0 22 

 †O. sp. 7 ?S---M, P ?P --- --- 18 0 0 0 18 

O. sp. 8 ?S---M, P ?P --- --- 4 0 0 0 4 

O. sp. 9 ?S---M, P ?P --- --- 2 0 0 0 2 

O. sp. 10 ?S---M, P ?P --- --- 0 0 1 5 6 

O. sp. 11 ?S---M, P ?P --- --- 1 0 0 1 2 

O. (lvara) sp. ?S---M, P ?P --- --- 1 0 0 0 1 

Peristichia sp. ?S---M, P ?P --- --- 1 0 0 0 1 

 Pyramidella sp. ?S---M, P ?P --- --- --- --- * --- * 

Turbonilla cf. pilsbryi (Bush, 1899) ?S---M, P ?P --- --- 0 0 0 1 1 

 †T. sp. 1 ?S---M, P ?P --- --- 0 0 9 14 23 

 †T. sp. 2 ?S---M, P ?P --- --- 0 0 0 15 15 

 T. sp. 3 ?S---M, P ?P --- --- 2 0 1 1 4 

 T. sp. 4 ?S---M, P ?P --- --- 0 0 0 1 1 

 T. sp. 5 ?S---M, P ?P --- --- 0 0 0 3 3 

 T. sp. 6 ?S---M, P ?P --- --- 3 0 1 0 4 

T. sp. ?S---M, P ?P --- --- 1 0 0 2 3 

Epitoniidae 

Epitonium sp. ?S---A ?P --- --- 0 0 0 1 1 

Eulimidae 

Melanella cf. arcuata (C.B. Adams, 1850) S---E P --- --- 1 0 0 5 6 

 †M.sp.1 S---E P --- --- 6 0 0 19 25 

 M. sp. S---E P --- --- 3 0 0 0 3 
Table B.2. (Continued). 

Appendices B-16 

MOE* FOOD† ZOOG‡ Z§ 6 m 27 m 42 m 80 m Total 

Calyptraeidae 

 †Crepidula aculeata (Gmelin, 1791) E F TC A/B 12 0 3 0 15 

C. fornicata (Linne, 1758) E F TN6 A 0 0 2 0 2 

†C. cf. fornicata (Linne, 1758) E F --- --- 2 0 9 1 12 

 C. plana (Say, 1822) E F TN6 A 0 1 0 0 1 

 C. sp. E F --- --- 0 1 0 1 2 

Capulidae 

Capulus cf. incurvatus (Gmelin, 1791) S---M F --- --- 0 0 0 1 1 

Cypraeidae 

Cypraea sp. F C/H --- --- --- --- * --- * 

Triviidae 

Erato maugeriae (Gray, 1832) S---T C TC A/B --- --- * --- * 

Trivia maltbiana (Schwengel and McGinty, 1942) S---T C TC A/B 0 0 3 0 3 

Velutinidae 

Lamellaria leucosphaera (Schwengel, 1942) F C T A 1 0 0 0 1 

L. perspicua (Linne, 1758) F C T A/B --- --- * --- * 

L. sp. F C --- --- 1 0 0 0 1 

Marsenina sp. F C --- --- 0 0 1 0 1 

?M. sp. F C --- --- 0 0 0 2 2 

Naticidae 

†Haliotinella patinaria (Guppy, 1876) F C T A 0 0 0 12 12 

Cymatiidae 

Cymatium sp. F C --- --- --- --- * --- * 

Order Neogastropoda 

Muricidae 

Aspella senex (Dall, 1903) F C TC --- --- --- * --- * 

Favartia cellulosa (Conrad, 1846) F C TC A --- --- * --- * 

Murexiella levicula (Dall, 1889) F C TC A 0 0 0 3 3 

M. macgintyi (M. Smith, 1938) F C T A/B 0 0 1 0 1 

Thais haemastoma floridana (Conrad, 1837) F C TC A --- --- * * 

Trachypollia didyma (Schwengel, 1943) F C TC A/B 0 0 0 1 1 

T. sp. F C --- --- 0 0 0 1 1 

 Coralliophilidae 

Coralliophila cf. aberrans (C.B. Adams, 1850)║ S---C O --- --- --- --- * --- * 

 C. sp. 1 S---C O --- --- 0 4 3 0 7 

 C. sp. 2║ S---C O --- --- 0 0 0 9 9 

 C. Sp. S---C 0 --- --- 0 0 1 0 1 

 †Latiaxis mansfieldi (McGinty, 1940) S---C 0 T B 0 0 2 15 17 

Buccinidae 

Antillophos adelus (Schwengel, 1942) F C/S T --- 0 0 0 7 7 

Colubraria lanceolata (Menke, 1828) F C/S TC A 0 0 0 1 1 

 C. swifti (Tryon, 1881) F C/S T A --- --- * --- * 

Engina corinnae (Crovo, 1971) F C/S T A 0 0 4 1 5 

E. turbinella (Kiener, 1835) F C/S T A 1 0 0 0 1 

 †Pisania tincta (Conrad, 1846) F C/S TC A 8 4 3 7 22 

Columbellidae 

 †Astyris lunata (Say, 1826) F C TN3 A 3 0 10 1 14 

Costoanachis floridana (Rehder, 1939) F C TC A 4 0 0 0 4 

 C. ?hotesseriana (Orbigny, 1842) F C --- --- 0 0 0 1 1 

 *C. lafresnayi (Fischer and Bernardi, 1856) F C TN5 B 0 0 34 585 619 

 *C. sparsa (Reeve, 1859) F C T A/B 14 2 87 55 158 

Costoanachis sp. F C --- --- 0 0 4 0 4 

 †Nassarina glypta (Bush, 1885) F C TC A/B 0 2 9 36 47 

Steironepion minor (C. B. Adams, 1845) F C TC A/B/C --- --- * --- * 

 S. monilifera (Sowerby, 1844) F C T A/B 1 0 0 0 1 

Suturoglypta iontha (Ravenel, 1861) F C TC A/B 0 0 3 0 3 

Nassariidae 

Nassarius consensus (Ravenel, 1861) F C/S TC A 0 0 2 1 3 

N. cf. consensus (Ravenel, 1861) F C/S --- --- 0 0 0 7 7 

N. sp. F C/S --- --- 0 0 2 0 2 

Fasciolariidae 

Fasciolaria tulipa (Linne, 1758) F C TC A --- --- * --- * 

Latirus cariniferus (Lamarck, 1822) F C T A --- --- * --- * 

 cf. Pleuroploca gigantea (Kiener, 1840) F C TC A --- --- * --- * 

Marginellidae 

Marginella lavalleeana (Orbigny, 1842) F ?C/S TC A/B 2 0 0 0 2 

Volvarina avena (Kiener, 1834) F C/S TC A --- --- --- * * 

 Marginellidae sp. F ?C/S --- --- 0 1 0 0 1 
Table B.2. (Continued). 

Appendices B-17 

MOE* FOOD† ZOOG‡ Z§ 6 m 27 m 42 m 80 m Total 

Costellariidae 

Pusia sykesi (Melvill, 1925) F C T A/B * * 

Turridae 

Cryoturris fargoi (McGinty, 1955) F C TC A/B --- --- --- * * 

Daphnella morra (Dall, 1881) F C TC A/B/C 0 0 0 1 1 

Glyphoturris rugirima (Dall, 1889) F C TC A/B/C 0 0 0 3 3 

lthycythara auberiana (Orbigny, 1842) F C T --- 0 0 2 0 2 

Nannodiella melanitica (Bush, 1885) F C TC A/B --- --- * --- * 

Pilsbryspira albomaculala (Orbigny, 1842) F C T A --- --- * --- * 

P. jayana (C.B. Adams, 1850) F C --- --- --- --- * --- * 

P. monilis (Bartsch and Rehder, 1939) F C T A 2 0 0 0 2 

Splendrillia sp. F C --- --- --- --- * --- * 

Subclass Opisthobranchia 

 Opisthobranchia sp. I F --- --- --- 0 0 1 0 1 

 Opisthobranchia sp. 2 F --- --- --- 0 0 0 1 1 

 Opisthobranchia sp. 3 F --- --- --- 0 0 1 0 1 

Order Cephalaspidea 

 Atyidae 

Atys caribaea (Orbigny, 1841) F ?C TC A/B 0 0 0 1 1 

Haminoea sp. F C --- --- 1 0 0 0 1 

Retusidae 

Pyrunculus caelatus (Bush, 1885) F ?C TC A/B 0 0 1 0 1 

Order Ascoglossa 

 Ascoglossa sp. F H --- --- 1 0 0 0 1 

Order Notaspidea 

 Pleurobranchidae 

 Pleurobrachidae sp. F C --- --- 3 0 0 0 3 

Order Thecosomata 

 Cavoliniidae 

Creseis acicula (Rang, 1828) F F TN P 0 0 0 1 1 

Order Nudibranchia 

 Goniodorididae 

?Okenia sp. F C * * 

 Polyceratidae 

?Polycera Sp. F C --- --- 0 0 0 6 6 

 Dotoidae 

Doto sp. ?S---H ?P --- --- 1 0 0 0 1 

 Aeolidiidae 

Spurilla neapolitana ?var. braziliana

(MacFarland, 1909) F C T --- 0 0 0 1 1 

 Gastropoda sp. 1 F --- --- --- 0 0 0 1 1 

 Gastropoda sp. 3 F --- --- --- 0 0 1 1 2 

 Gastropoda sp. F --- --- --- 0 2 1 1 4 

Class Scaphopoda 

 Order Dentaliida 

 Dentaliidae 

Dentalium sp. F F --- --- --- --- * --- * 

 Order Gadilida 

 Siphonodentaliidae 

Cadulus sp. F F --- --- 0 0 0 2 2 

Class Polyplacophora 

 Order Ischnochitonida 

 Ischnochitonidae 

Ischnochiton sp. F H --- --- 1 0 0 0 1 

Callistoplacidae 

Callistochiton sp. F H --- --- 0 0 0 1 1 

Chaetopleuridae 

Chaetopleura apiculala (Say, 1830) F H TN3 A 0 0 0 6 6 

 C. cf. apiculata (Say, 1830) F H --- --- 0 0 0 7 7 

 Order Acanthochitonida 

 Acanthochitonidae 

Acanthochitona sp. F H --- --- 1 0 0 1 2 

 Class Bivalvia 

 Order Nuculoida 

 Nuculidae 

Nucula ?aegensis (Jeffreys, 1879) F D --- --- 0 0 0 1 1 

N. crenulata (A. Adams, 1856) F D TC A 0 0 0 4 4 
Table B.2. (Continued). 

Appendices B-18 

MOE* FOOD† ZOOG‡ Z§ 6 m 27 m 42 m 80 m Total 

 Nuculanidae 

Portlandia minuscula (Verrill and Bush, 1897) F D M3 C 0 0 0 3 3 

 Order Arcoida 

 Arcidae 

 Anadara floridana (Conrad, 1869) E F TC A 0 0 0 1 1 

†Arca imbricata Bruguiere, 1789 E F TC A 35 2 1 0 38 

A. zebra (Swainson, 1833) E F TC A 0 1 0 1 2 

Arcopsis adamsi (Dall, 1886) E F TC A 0 0 0 7 7 

Barbatia cancellaria (Lamarck, 1819) E F TC A 0 1 0 0 1 

 *B. candida (Helbling, 1779) E F TC A 1 1 8 157 167 

 †B. domingensis (Lamarck, 1819) E F TC A 16 0 0 0 16 

 Order Mytiloida 

 Mytilidae 

 †Gregariella coralliophaga (Gmelin, 1791) N F TC A 6 4 8 81 99 

 Lioberus castaneus (Say, 1822) E F T A --- --- * --- * 

 †Lithophaga aristata (Dillwyn, 1817) N F TC A/B/C 3 7 18 0 28 

 *L. bisulcata (Orbigny, 1842) N F TC A/B 150 6 94 17 267 

 Modiolus cf. modiolus squamosus (Beauperthuy, 

1967) E F --- --- --- --- * --- * 

 †Musculus lateralis (Say, 1822) E F TC A 8 2 3 1 14 

Mytilidae sp. E F --- --- 0 0 0 2 2 

 Order Pterioida 

 Pteriidae 

Pteria colymbus (Roding, 1798) E F TC A 1 0 0 3 4 

 Isognomonidae 

lsognomon bicolor (C.B. Adams, 1845) E F T A 1 0 0 0 1 

 Order Limoida 

 Limidae 

Lima lima (Linne, 1758) E F TC A/B 1 0 0 0 1 

 Order Ostreoida 

 Ostreidae 

 Ostrea permollis (Sowerby, 1841) E F TC A 0 0 0 2 2 

 †Ostreola equestris (Say, 1834) E F TN1 A 18 0 0 6 24 

 †Ostreidae sp. E F --- --- 0 0 0 13 13 

 Gryphaeidae 

*Neopycnodonte cochlear (Poli, 1795) E F TC A/B/C 2 0 3 161 166 

 Plicatulidae 

 †Plicatula gibbosa (Lamarck, 1801) E F TC A 0 0 0 18 18 

 Pectinidae 

*Chlamys benedicti (Verrill and Bush, 1897) E F TC A 0 0 26 93 119 

Lyropecten nodosus (Linne, 1758) E F TC A --- --- * * 

 Pectinidae sp. E F --- --- 0 0 0 1 1 

 Anomiidae 

Anomia simplex (Orbigny, 1842) E F TN3 A 1 0 0 1 2 

Pododesmus rudis (Broderip, 1834) E F TC A 2 0 1 4 7 

 Order Hippuritoida 

 Chamidae 

 †Chama congregata (Conrad, 1833) E F TC A 1 4 10 2 17 

 C. florida (Lamarck, 1819) E F T A/B 0 1 0 1 2 

 C. lactuca (Dall, 1886) E F TC A 0 0 3 5 8 

 C. macerophylla (Gmelin, 1791) E F TC A 0 1 0 1 2 

 C. sarda (Reeve, 1847) E F T B 0 1 0 0 1 

 C. sp. E F --- --- 0 0 0 1 1 

Pseudochama radians (Lamarck, 1819) E F TC A/B 2 0 0 0 2 

 Order Veneroida 

 Lucinidae 

 †Parvilucillu mulLilineata (Tuomey and Holmes, 

1857) F F TC A/B 0 0 0 14 14 

 Thyasiridae 

Thyasira flexuosa (Montagu, 1803) F F B7 A/B 0 0 0 1 1 

 Ungulinidae 

Diplodonta punctata (Say, 1822) F F TC A/B/C --- --- --- * * 

Diplodonta sp. F F --- --- 2 0 0 2 4 

 Leptonidae 

Montacuta sp. S---E F --- --- 0 0 0 1 1 

 Leptonidae sp. ?S---E F --- --- 1 0 1 2 4 
Table B.2. (Continued). 

Appendices B-19 

MOE* FOOD† ZOOG‡ Z§ 6 m 27 m 42 m 80 m Total 

 Lasaeidae 

Aligena cf. texasiana (Harry, 1969) ?S---E, P F --- --- 1 0 0 0 1 

A. sp. ?S---E, P F --- --- 1 0 0 0 1 

Erycina sp. ?S---? F --- --- 0 0 0 6 6 

Kellia suborbicularis (Montagu, 1803) F F B A 0 0 2 3 5 

K. ?suborbicularis (Montagu, 1803) F? F --- --- 2 0 0 0 2 

 †K. sp. ?S---? F --- --- 5 0 2 4 11 

 Lasaeidae sp. ?S---? F --- --- 1 0 0 6 7 

 Condylocardiidae 

Carditopsis smithii (Dall, 1896) F F T A 2 0 0 0 2 

 Crassatellidac 

Crassinella lunulata (Conrad, 1834) F F TN3 A/B 0 0 3 1 4 

 Cardiidae 

Cerastoderma cf. pinnatulatum (Conrad, 1831) F F --- --- 1 0 0 0 1 

 Tellinidae 

Tellina cf. squamifera (Deshayes, 1855) F D --- --- 1 0 0 0 1 

 †T. cf. sybaritica (Dall, 1881) F D --- --- 2 0 5 6 13 

T. sp. F D --- --- 1 0 1 0 2 

 Vesicomyidae 

Calyptogena sp. F F --- C? 1 0 0 0 1 

 Veneridae 

*Chione grus (Holmes, 1858) F F TC A 110 8 30 1 149 

Cyclinella tenuis (Recluz, 1852) F F TN1 A/B 7 0 0 0 7 

 Veneridae sp. F F --- --- 0 0 3 2 5 

 Petricolidae 

Rupellaria typica (Jonas, 1844) N F TC A 2 0 0 0 2 

 Order Myoida 

 Myidae 

Sphenia antillensis (Dall and Simpson, 1901) E F T A 9 0 0 0 9 

 Gastrochaenidae 

 †Gastrochaena hians (Gmelin, 1791) N F TC --- 1 0 4 41 46 

 †Rocellaria ovata (Sowerby, 1834) N F TC --- 3 0 6 12 21 

 Hiatellidae 

 Hiatella arctica (Linne, 1767) E F B A/B/C 0 0 2 7 9 

 Pholadidae 

 *Dip/othyra smithii (Tryon, 1862) N F TN3 A 207 0 0 0 207 

Subclass Anomalodesmata 

 Order Pholadomyoida 

 Lyonsiidae 

 Lyonsia beana (Orbigny, 1842) E F TC A 3 0 4 0 7 

 Bivalvia sp. 2 --- 1 0 0 0 1 

 Bivalvia sp. 4 --- 1 1 0 4 6 

 Bivalvia sp. 7 --- 0 0 0 1 1 

 Bivalvia sp. 8 --- 0 0 0 1 1 

 Bivalvia sp. --- 3 2 1 3 9 

Class Cephalopoda 

 Subclass Coleoidea 

 Order Teuthoidea 

 Ommastrephidae 

 lllex oxygoneus (Roper, Li and Mangold, 1969) F C TN3 B/C --- --- --- * * 

Order Octopoda 

 Octopodidae 

 Octopus sp. F C --- --- 0 1 0 0 1 

* Mode of existence (MOE): E = epilithic; F = free-living; N - endolithic; S = symbiotic (-A = with anthozoan; -C = with scleractinian coral; E = 

with echinoderm: -H = with hydroid; -M = with mollusk; -P = with polychaete; -S = with sponge; -T = with tunicate [ascidian]). 

† Feeding type (FOOD): C = carnivore; D = detritivore/deposit-feeder; F = filter-feeder/suspension-feeder/mucoid entrapper; H = herbivore; 

0 = corallivore; P = parasite; S = scavenger. 

‡ Zoogeographical distribulion (ZOOG): B = Areticlboreal; M - temperate; T = tropicaVstenothermic; TC = IropicaVto Cape Hatteras; TN = 

tropical/to nonh of Cape Hatteras [I = to Virginia; 2 - to Maryland; 3 = to Massachusetts; 4 = 10 New Hampshire; 5 = to Maine; 6 = to 

Canada; 7 = to Greenland]. 

§ Depth (Z): A = shallow (0-50 m); B = shelf (50-200 m); C - deep (>200 m); P - pelagic. 

║ FOOD catcagorization of these species reflects the results of gut analyses in the present study. 

# Odostomia is placed in quotes according to Robertson's (1978) findings that odostomoid species require biological data (e.g., on 

spermatophores, reproductive anatomy insufficient here for proper generic placement). 
Appendices B-20 

Table B.3. Decapod crustacean community composition and total number of individuals per species on 

east Florida Oculina coral reefs (From: Reed et al., 1982). 

6 m 

Fort Pierce Inlet 

27 m 

North of 

Fort Pierce 

42 m 

South of 

Fort Pierce 

St. Lucie Inlet 

80 m 

Offshore East of

Fort Pierce 

Families 11 9 10 10 

Genera 21 13 13 20 

Species 30 14 16 24 

Endemic Species 16 1 3 9 

Mean Species 7.9 6.8 5.9 10.9 

Mean Individuals 39.8 18.5 47 87.6 

Mean Coral Size (g) 521.8 131.4 1,193.50 1,613.60 

Percent Dead Coral 23.3 18.3 21.9 69.9 

Number Samples 15 4 13 10 

Dominance (D1) 40 32 72 18 

Dominance (D2) 55 45 79 34 

Species 

1. Pagurus carolinensis 6 24 439 160 

2. Megalobrachium soriatum 238 9 41 22 

3. Pagurus piercei 0 0 37 142 

4. Synalpheus townsendi 1 8 31 112 

5. Galathea rastrata 0 1 5 142 

6. Micropanope scuptipes 1 1 0 137 

7. Pachycheles monilifer 89 0 0 2 

8. Mithrax forceps 81 8 0 0 

9. Thor manningi 49 3 0 29 

10. Pseudomedaeus distinctus 0 0 11 33 

11. Synalpheus fritzmuelleri 30 0 0 0 

12. Pseudomedaeus agassizii 0 0 10 19 

13. Nematopaguroides pusillus 0 0 0 29 

14. Periclimenes iridescens 2 1 25 0 

15. Synalpheus minus 19 0 0 0 

16. Micropanope barbadensis 0 0 0 17 

17. Synalpheus cf. townsendi 15 0 1 0 

18. Pelia mutica 4 9 1 1 

19. Euchirograpsus americanus 1 0 0 10 

20. Stenorhynchus seticornis 0 3 3 5 

21. Pilumnus dasypodus 10 0 0 0 

22. Petrolisthes galathinus 9 0 0 0 

23. Periclimenes rathbunae 8 0 0 0 

24. Troglocarcinus corallicola 6 0 1 0 

25. Periclimenaeus atlanticus 6 0 0 0 

26. Domecia acanthophora 5 0 0 0 

27. Micropanope nuttingi 0 3 0 2 

28. Mithrax acuticornis 0 0 1 3 

29. Paguristes tortugae 2 2 0 0 

30. Nanoplax xanthiformis 0 0 0 4 

31. Lobopilumnus agassizii 2 1 0 0 
Table B.3. (Continued). 

Appendices B-21 

6 m 

Fort Pierce Inlet 

27 m 

North of 

Fort Pierce 

42 m 

South of 

Fort Pierce 

St. Lucie Inlet 

80 m 

Offshore East of

Fort Pierce 

32. Synalpheus cf. fritzmuelleri 3 0 0 0 

33. Alpheus cristulifrons 2 0 0 0 

34. Gonodactylus bredini 2 0 0 0 

35. Scyllarus depressus 0 0 2 0 

36. Pinnixa retinens 0 0 2 0 

37. Parapinnixa hendersoni 0 0 0 2 

38. Porcellana sigsbeiana 1 0 0 0 

39. Periclimenes americanus 1 0 0 0 

40. Epialtus sp. 1 0 0 0 

41. Alpheus formosus 1 0 0 0 

42. Eucratopsis crassimanus 1 0 0 0 

43. Thor dobkini 1 0 0 0 

44. Paractaea rufopunctata nodosa 0 1 0 0 

45. Stenocionops furcata 0 0 1 0 

46. Periclimenes longicaudatus 0 0 0 1 

47. Solenopagurus lineatus 0 0 0 1 

48. Automate evermanni 0 0 0 1 

49. Aplheus cf. amblyonyx 0 0 0 1 

50. Pinnotheres sp. 0 0 0 1 
Appendices C-1 

Appendix C
Appendices C-2 Table C.1. Fish species recorded from natural nearshore hardbottom habitats in southeast and east central Florida. 68 Families; 257 spp. Trophic Category** Residency Size Classes 

Family Species Orig. Cite* H Pl D I P Res Tran Onto <2 2 to 10 10 to 25 25 to 50 50 to 100 >100 NS 

GINGLYMOSTOMATIDAE Ginglymostoma cirratum 1 ● ● ● 

Carcharhinus brevipinna 3 ● ● ● Carcharhinus leucas 1 ● ● ● Carcharhinus limbatus 1 ● ● ● 

CARCHARHINIDAE Carcharhinus plumbeus 1 ● ● ● SPHYRNIDAE Sphyrna mokarran 6 ● ● ● RHINOBATIDAE Rhinobatos lentiginosus 5 ● ● ● DASYATIDAE Dasyatis americana 4 ● ● ● MYLIOBATIDAE Aetobatus narinari 4 ● ● ● UROLOPHIDAE Urobatis jamaicensis 4 ● ● ● MOBULIDAE Manta birostris 4 ● ● ● ELOPIDAE Megalops atlanticus 3 ● ● ● 

Echidna catenata 3 ● ● ● Enchelycore carychroa 2 ● ● Gymnothorax funebris 1 ● ● ● Gymnothorax moringa 1 ● ● ● Gymnothorax vicinus 4 ● ● ● 

MURAENIDAE Muraena miliaris 3 ● ● ● 

Ahlia egmontis 2 ● ● ● 

OPHICHTHIDAE Myrichthys breviceps 4 ● ● ● 

Myrichthys ocellatus 5 ● ● ● Harengula clupeola 1 ● ● ● Harengula humeralis 1 ● ● ● Harengula jaguana 1 ● ● ● Opisthonema oglinum 1 ● ● ● 

CLUPEIDAE Sardinella aurita 1 ● ● ● 

Anchoa cubana 1 ● ● ● 

ENGRAULIDAE Anchoa hepsetus 1 ● ● ● 

Anchoa lyolepis 1 ● ● ● Synodus foetens 4 ● ● ● ● 

SYNODONTIDAE Synodus intermedius 4 ● ● ● GOBIESOCIDAE Gobiesox strumosus 1 ? ● ● 

Membras martinica 1 ● ● ● 

ATHERINIDAE Menidia peninsulae 1 ● ● ● 
Table C.1. (Continued).

Appendices C-3 68 Families; 257 spp. Trophic Category** Residency Size Classes 

Family Species Orig. Cite* H Pl D I P Res Tran Onto <2 2 to 10 10 to 25 25 to 50 50 to 100 >100 NS 

BELONIDAE Tylosurus crocodilus 5 ● ● ● ● ● 

Hemiramphus brasiliensis 3 ● ● ● 

HEMIRAMPHIDAE Hyporhamphus unifasciatus 1 ● ● ● 

Holocentrus adscensionis 4 ● ● ● ● ● Holocentrus rufus 3 ● ● ● 

HOLOCENTRIDAE Myripristis jacobus 4 ● ● ● ● ● AULOSTOMIDAE Aulostomus maculatus 1 ● ● ● ● ● FISTULARIIDAE Fistularia tabacaria 4 ● ● ● 

Cosmocampus albirostris 4 ● ● ● Cosmocampus elucens 4 ● ● ● 

SYNGNATHIDAE Scorpaena grandicornis 5 ● ● ● ● ● SCORPAENIDAE Scorpaena plumieri 1 ● ● ● ● ● TRIGLIDAE Prionotus scitulus 4 ● ● ● DACTYLOPTERIDAE Dactylopterus volitans 4 ● ● ● CENTROPOMIDAE Centropomus undecimalis 3 ● ● ● ● 

Centropristis striata 1 ● ● ● ● ● Cephalopholis fulva 5 ● ● ● ● ● Cephalopholis cruentata 4 ● ● ● ● ● Diplectrum formosum 4 ● ● ● ● ● Epinephelus adscensionis 4 ● ● ● ● ● Epinephelus itajara 1 ● ● ● ● ● Epinephelus morio 1 ● ● ● ● ● Hypoplectrus puella 4 ● ● ● Hypoplectrus unicolor 4 ● ● ● Mycteroperca bonaci 3 ● ● ● Mycteroperca microlepis 1 ● ● ● Mycteroperca phenax 4 ● ● ● Rypticus maculatus 1 ● Rypticus saponaceus 4 ● Rypticus subbifrenatus 6 ● Serranus baldwini 4 ● ● ● ● Serranus subligarius 1 ● ● ● ● 

SERRANIDAE Serranus tigrinus 4 ● ● ● ● ● 
Table C.1. (Continued).

Appendices C-4 68 Families; 257 spp. Trophic Category** Residency Size Classes 

Family Species Orig. Cite* H Pl D I P Res Tran Onto <2 2 to 10 10 to 25 25 to 50 50 to 100 >100 NS 

Apogon binotatus 1 ● ● ● ● Apogon maculatus 1 ● ● ● ● Apogon pseudomaculatus 1 ● ● ● ● Apogon townsendi 4 ● ● ● ● Astrapogon puncticulatus 4 ● ● ● ● Astrapogon stellatus 2 ● ● ● ● 

APOGONIDAE Phaeoptyx conklini 2 ● ● ● ● 

Caranx bartholomaei 1 ● ● ● ● ● ● ● Caranx crysos 1 ● ● ● ● ● ● Caranx hippos 1 ● ● ● ● ● ● Caranx latus 1 ● ● ● ● ● ● Caranx ruber 1 ● ● ● ● ● ● Chloroscombrus chrysurus 1 ● ● ● ● Decapterus punctatus 3 ● ● ● ● ● ● Elagatis bipinnulata 4 ● ● ● Oligoplites saurus 1 ● ● ● ● Selar crumenophthalmus 3 ● ● ● ● ● ● Selene setapinnis 1 ● ● ● ● ● Selene vomer 1 ● ● ● ● ● Seriola dumerili 4 ● ● ● ● ● Seriola rivoliana 5 ● ● ● ● ● Trachinotus carolinus 3 ● ● ● ● ● Trachinotus falcatus 3 ● ● ● ● 

CARANGIDAE Trachinotus goodei 6 ● ● ● 

Lutjanus apodus 1 ● ● ● ● ● Lutjanus analis 1 ● ● ● ● ● ● Lutjanus griseus 1 ● ● ● ● ● ● Lutjanus jocu 1 ● ● ● ● Lutjanus mahogoni 1 ● ● ● ● ● Lutjanus synagris 1 ● ● ● ● ● Ocyurus chrysurus 1 ● ● ● ● 

LUTJANIDAE Rhomboplites aurorubens 6 ● ● ● ● LOBOTIDAE Lobotes surinamensis 5 ● ● ● 
Table C.1. (Continued).

Appendices C-5 68 Families; 257 spp. Trophic Category** Residency Size Classes 

Family Species Orig. Cite* H Pl D I P Res Tran Onto <2 2 to 10 10 to 25 25 to 50 50 to 100 >100 NS 

Eucinostomus argenteus 1 ● ● ● ● Eucinostomus gula 1 ● ● ● ● Eucinostomus jonesii 4 ● ● ● ● Eucinostomus lefroyi 4 ● ● ● Eucinostomus melanopterus 6 ● ● ● ● 

GERREIDAE Gerres cinereus 1 ● ● ● ● ● 

Anisotremus surinamensis 1 ● ● ● ● ● Anisotremus virginicus 1 ● ● ● ● ● Haemulon album 4 ● ● ● ● ● Haemulon aurolineatum 1 ● ● ● ● ● Haemulon carbonarium 1 ● ● ● ● ● Haemulon chrysargyreum 1 ● ● ● ● ● ● Haemulon flavolineatum 1 ● ● ● ● ● Haemulon macrostomum 3 ● ● ● ● ● Haemulon melanurum 1 ● ● ● ● ● Haemulon parra 1 ● ● ● ● ● Haemulon plumierii 1 ● ● ● ● ● Haemulon sciurus 3 ● ● ● ● ● Haemulon striatum 4 ● ● ● ● 

HAEMULIDAE Orthopristis chrysoptera 5 ● ● ● ● ● 

Archosargus probatocephalus 1 ● ● ● Archosargus rhomboidalis 4 ● ● ● ● Calamus bajonado 1 ● ● ● ● Calamus calamus 4 ● ● ● ● ● ● Calamus penna 4 ● ● ● ● ● ● Diplodus argenteus 1 ● ● ● ● ● ● ● ● ● Diplodus holbrooki 1 ● ● ● ● ● ● ● ● ● 

SPARIDAE Lagodon rhomboides 5 ● ● ● POLYNEMIDAE Polydactylus virginicus 6 ●
Table C.1. (Continued).

Appendices C-6 68 Families; 257 spp. Trophic Category** Residency Size Classes 

Family Species Orig. Cite* H Pl D I P Res Tran Onto <2 2 to 10 10 to 25 25 to 50 50 to 100 >100 NS 

Bairdiella sanctaeluciae 1 ● ● ● ● ● ● Cynoscion nebulosus 6 ● ● ● ● Equetus acuminatus 1 ● ● ● Equetus umbrosus 1 ● ● ● Sciaenops ocellatus 6 ● ● ● ● ● Odontoscion dentex 1 ● ● ● 

SCIAENIDAE Umbrina coroides 3 ● ● 

Mulloidichthys martinicus 3 ● ● ● ● 

MULLIDAE Pseudupeneus maculatus 1 ● ● ● ● PEMPHERIDAE Pempheris schomburgkii 1 ● ● ● ● KYPHOSIDAE Kyphosus sectatrix/incisor 1 ● ● ● ● ● EPHIPPIDAE Chaetodipterus faber 1 ● ● ● ● ● 

Chaetodon capistratus 5 ● ● ● ● Chaetodon ocellatus 3 ● ● ● Chaetodon sedentarius 4 ● ● ● ● 

CHAETODONTIDAE Chaetodon striatus 5 ● ● ● 

Holacanthus bermudensis 3 ● ● ● ● ● ● ● Holacanthus ciliaris 1 ● ● ● ● ● ● ● Pomacanthus arcuatus 1 ● ● ● ● ● ● ● 

POMACANTHIDAE Pomacanthus paru 2 ● ● ● ● ● ● ● 

Abudefduf saxatilis 1 ● ● ● ● ● ● Abudefduf taurus 1 ● ● ● ● ● ● Microspathodon chrysurus 4 ● ● ● ● Stegastes adustus 5 ● ● ● ● ● Stegastes diencaeus 4 ● ● ● ● ● Stegastes leucostictus 1 ● ● ● ● ● Stegastes partitus 2 ● ● ● ● ● ● Stegastes planifrons 4 ● ● ● 

POMACENTRIDAE Stegastes variabilis 1 ● ● ● ● 

Opistognathus aurifrons 5 ● ● ● Opistognathus macrognathus OPISTOGNATHIDAE 1 ● ● ● Opistognathus whitehursti 4 ● ● ● 
Table C.1. (Continued).

Appendices C-7 68 Families; 257 spp. Trophic Category** Residency Size Classes 

Family Species Orig. Cite* H Pl D I P Res Tran Onto <2 2 to 10 10 to 25 25 to 50 50 to 100 >100 NS 

Mugil cephalus 3 ● ● ● 

MUGILIDAE Mugil curema 3 ● ● ● 

Bodianus rufus 3 ● ● ● ● ● Doratonotus megalepis 1 ● ● ● ● Halichoeres bivittatus 1 ● ● ● ● ● Halichoeres cyanocephalus 4 ● ● ● ● Halichoeres garnoti 4 ● ● ● ● Halichoeres maculipinna 1 ● ● ● ● ● Halichoeres poeyi 1 ● ● ● ● ● Halichoeres radiatus 1 ● ● ● ● ● Lachnolaimus maximus 3 ● ● ● Thalassoma bifasciatum 1 ● ● ● ● ● Xyrichtys martinicensis 4 ● ● ● ● ● 

LABRIDAE Xyrichtys splendens 4 ● ● ● ● ● 

Cryptotomus roseus 4 ● Scarus coelestinus 1 ● ● ● ● ● ● Scarus coeruleus 4 ● ● ● ● ● ● Scarus guacamaia 1 ● ● ● ● ● ● Scarus iseri 5 ● ● ● ● ● Scarus taeniopterus 4 ● ● ● ● ● Scarus vetula 3 ● ● ● ● ● ● Sparisoma atomarium 1 ● ● ● ● Sparisoma aurofrenatum 3 ● ● ● ● ● Sparisoma chrysopterum 3 ● ● ● ● ● Sparisoma radians 4 ● ● ● ● Sparisoma rubripinne 1 ● ● ● ● ● ● 

SCARIDAE Sparisoma viride 3 ● ● ● ● ● ● URANOSCOPIDAE Astroscopus y-graecum 1 ● ● ● TRIPTERYGIIDAE Enneanectes pectoralis 5 ● ● ● ●
Table C.1. (Continued).

Appendices C-8 68 Families; 257 spp. Trophic Category** Residency Size Classes 

Family Species Orig. Cite* H Pl D I P Res Tran Onto <2 2 to 10 10 to 25 25 to 50 50 to 100 >100 NS 

Labrisomus bucciferus 3 ● ● ● ● Labrisomus gobio 1 ● ● ● ● Labrisomus kalisherae 4 ● ● ● ● Labrisomus nuchipinnis 1 ● ● ● ● ● Malacoctenus aurolineatus 5 ● ● ● ● Malacoctenus macropus 1 ● ● ● ● Malacoctenus triangulatus 1 ● ● Paraclinus fasciatus 4 ● Paraclinus marmoratus 4 ● Paraclinus nigripinnis 1 ● LABRISOMIDAE Starksia ocellata 1 ● Acanthemblemaria aspera 4 ● ● ● ● Acanthemblemaria spinosa CHAENOPSIDAE 4 ● ● ● ● Emblemaria pandionis 4 ● ● ● ● Dactyloscopus crossotus 1 ● DACTYLOSCOPIDAE Platygillellus rubrocinctus 2 ● Entomacrodus nigricans 1 ● Hypleurochilus bermudensis 4 ● ● ● ● Ophioblennius macclurei 4 ● Parablennius marmoreus 2 ● ● ● ● 

BLENNIIDAE Scartella cristata 1 ● ● ● ● CALLIONYMIDAE Paradiplogrammus bairdi 5 ● ● ● ● ELEOTRIDAE Erotelis smaragdus 3 ● Coryphopterus dicrus 4 ● ● ● ● 

Coryphopterus eidolon 5 ● ● ● ● Coryphopterus glaucofraenum 3 ● ● ● ● Coryphopterus personatus 4 ● ● ● ● Ctenogobius saepepallens 4 ● ● ● Elacatinus macrodon 5 ● ● ● 

GOBIIDAE Elacatinus oceanops 5 ● ● ● ● 
Table C.1. (Continued).

Appendices C-9 68 Families; 257 spp. Trophic Category** Residency Size Classes 

Family Species Orig. Cite* H Pl D I P Res Tran Onto <2 2 to 10 10 to 25 25 to 50 50 to 100 >100 NS 

Gnatholepis thompsoni 4 ● ● ● ● Gobiosoma grosvenori 4 ● Microgobius carri 4 ● ● ● Nes longus 3 ● ● ● 

GOBIIDAE (continued) Priolepis hipoliti 4 ● Ptereleotris calliurus 4 ● PTERELEOTRIDAE Ptereleotris helenae 4 ● Acanthurus bahianus 1 ● ● ● ● ● ● ● ACANTHURIDAE Acanthurus chirurgus 1 ● ● ● ● ● ● ● 

Acanthurus coeruleus 1 ● ● ● ● ● ● ● Sphyraena barracuda 1 ● ● ● ● ● ● ● ● 

SPHYRAENIDAE Sphyraena guachancho 1 ● ● ● ● ● 

Sphyraena picudilla 5 ● ● ● ● ● Pomatomus saltatrix 1 ● ● ● 

POMATOMIDAE Scomberomorus maculatus 5 ● ● ● SCOMBRIDAE Scomberomorus regalis 3 ● ● ● ● BOTHIDAE Bothus lunatus 5 ● ● ● ● ● 

Balistes capriscus 4 ● ● ● ● ● ● 

BALISTIDAE Balistes vetula 4 ● ● ● ● 

Canthidermis sufflamen 5 ● ● ● ● ● ● Aluterus schoepfii 4 ● ● ● ● ● ● Aluterus scriptus 3 ● ● ● ● ● ● Cantherhines macrocerus 5 ● ● ● Cantherhines pullus 3 ● ● ● Monacanthus tuckeri 5 ● ● ● ● ● 

FILEFISHES Stephanolepis hispidus 1 ● ● ● 

Acanthostracion polygonia 4 ● ● ● Acanthostracion quadricornis 3 ● ● ● Lactophrys bicaudalis 4 ● ● ● Lactophrys trigonus 4 ● ● ● 

OSTRACIIDAE Lactophrys triqueter 1 ● ● ● 

Canthigaster rostrata 3 ● ● ● ● 

TETRAODONTIDAE Sphoeroides spengleri 2 ● ● ● 

Sphoeroides testudineus 5 ● ● ● ● 
Table C.1. (Continued).

Appendices C-10 68 Families; 257 spp. Trophic Category** Residency Size Classes 

Family Species Orig. Cite* H Pl D I P Res Tran Onto <2 2 to 10 10 to 25 25 to 50 50 to 100 >100 NS 

Chilomycterus schoepfii 4 ● ● ● 

DIODONTIDAE Diodon holocanthus 4 ● ● ● ● 

Diodon hystrix 3 ● ● ● ● 

Totals 31 36 2 123 74 136 57 104 130 166 100 41 18 14 0 

* Original Citation: 1 = Gilmore, 1977; Gilmore et al., 1981. 2 = Futch and Dwinell, 1977. 3 = Lindeman, 1997a; Lindeman and Snyder, 1999. 4 = Baron et al., 2004. 5 = Co

6 = Present study. ** H = Herbivore; Pl = Planktivore; D = Detritivore; I = Invertivore; P = Piscivore. 
Appendices D-1 

Appendix D
Appendices D-2 Table D.1. Examples of controlled experiments and field analyses of similar nearshore hardbottom genera/species inverte

Mesograzer Species Artificial Diet Foraging Items Location/ Species Ran

Littorina obtusata Ascophyllum nodosum Ascophyllum nodosum (ungrazed beforehand) North Atlantic Idotea granulose Ascophyllum nodosum Ascophyllum nodosum North Atlantic Placida dendritica (sea slug) --- Codium setchellii (Trowbridge, 1992) SE Australia Ampithoe valida Gracilaria (Nicotri, 1977) --- North Atlantic Idotea baltica Gracilaria (Nicotri, 1977) --- North Atlantic/ B

Sea 

Littorina littorea Fucus distichus (Van Alstyne, 1990) --- North Atlantic Hyale media F. vesiculosus --- Brazil Acmaea inessa --- Kelp: Egregia laevigata (Setchell) Brazil Scurria scurra --- Lessonia nigrescens (Munoz and Santelices, 1989) Brazil Hyale --- Iridaea laminarioides Brazil Aplysia californica Gracilaria ferox --- California Mytilus californiannus --- Odonthalia floccose Oregon rocky intertidal Diadema antillarum (urchin) Dictyota pulchella, Lobophora variegate --- --- Ampithoe longimana Ulva sp. (Enteromorpha), Hypnea musciformis, Gracilaria tikvahiae. Did not discriminate Red, brown, green algae. Diatoms (Cousfield, 1973; Nelson, 1979b; Hay, 1987, Duffy and Hay, 1991b, 1994) detritus Western Atlantic Elasmopus levis Ulva sp. (Enteromorpha), Hypnea musciformis, Gracilaria tikvahiae --- East coast Gammarus mucronatus Ulva sp. (Enteromorpha), Hypnea musciformis, Gracilaria tikvahiae. Gut content, detritus, diatoms epiphytic algae, macroalgae (Sanders, 1962; Zimmerman, 1979; Duffy and Hay, 1994) East coast Ampithoe marcuzii --- Sargassum filipendula North Carolina Caprella penantis --- Sargassum filipendula North Carolina Jassa falcate --- Sargassum filipendula North Carolina Erichthonius brasiliensis Sargassum filipendula --- North Carolina Elysia rufescens, ornata Bryopsis pennata --- Hong Kong Menaethius monoceros (crab) Acanthophora spicifera --- Hong Kong 
Table D.1. (Continued). 

Appendices D-3 Mesograzer Species Artificial Diet Foraging Items Location/ Species Ran

Ampithoe longimana --- Sargassum filipendula, Dictyota, Padina gymnospora, Hypnea musciformis, Chondria dasyphylla, Ulva North Carolina Ampithoe longimana Dictyota dichotoma, Calonitophyllum medium (foliose) Hypnea musciformis and Chondria dasyphylla, Sargassum filpendula --- North Carolina Limpets, coiled snails, chitons, isopods, amphipods, gastropod --- Halimeda opuntia, Amphiroa sp, Bryothamnion seaforthii occur on more coral and rock. Sandy areas had Gracilaria, Halymenia, Solieria. Herbivore abundance correlate with patterns. Oregon Ilyanassa obsolete Ulva lactuca --- Hog Island Bay, Virginia Littorina striata No-choice lab experiments - high feeding Ulva (Enteromorpha muscoides), Ulva rigida. Little to no feeding on Alsidium corallinum and Laurencia corrallopsis --- Gran Canaria (w

algae was collec

Osilinus atratus No-choice lab experiments - high feeding Ulva (Enteromorpha muscoides), Ulva rigida. Little to no feeding on Alsidium corallinum and Laurencia corrallopsis --- Gran Canaria (w

algae was collec

Grapsus albolineatus Filamentous algae Hong Kong Grapsus albolineatus Enteromorpha clathrata, Hincksia mitchelliae, Chaetomorpha antennina (filamentous which was eaten). Dermonema frappieri, Pterocladia tenuis, Porphyra suborbiculata, Ulva fasciata, and Endarachne Binghamiae (foliose which wasn’t eaten) --- Hong Kong Viviparus viviparous Chatophora incrassata, Chateophora pisiformis, Cladophora, Coleochaete pulvinata, Coleochaete scutata, mougeoutia, Oedogonium, Spirogyra tenuissima, Cyanos Anabaena, and different families of Rivulariacea, cyanophyta (12), Bacillariophyceae --- Sweden 
Table D.1. (Continued). 

Appendices D-4 Mesograzer Species Artificial Diet Foraging Items Location/ Species Ran

Littorina brevicula and Haliotis discus Ecklonia cava --- Sangju, South KDiadema antillarum Philippi Halimeda opuntia, Lobophora variegata, Sargassum sp., Galaxaura sp. --- Discovery Bay, Jamaica Acmaea testudinalis Clathromorphum circumscription --- Subarctic weste

North Atlantic 

Molluscs (divided by radulae-type) Cladophora, Polysiphonia, Ulva, Porphyra, Chondrus, Gigartina, Laminaria, Fucus, Corallina, Bossiella, Lithothamnium, Clathromorphum --- Maryland Sea urchins S. franciscanus, S. drobachiensis, S. purpuratus Nereocystis (liked), Costaria, L. saccharina, Callophyllis flabellulata, M. fuscum, A. cribrosum A. fimbriatum (disliked) --- San Juan Island 

Washington coa

Strongylocentrotus purpuratus (urchin), Lacuna porrecta, Tegula funebralis (snails), Idotea wosnesenskii (isopod) Kelps Brown algae: Fucus gardneri and F. spiralis --- Boiler Bay, OregAmpithoe ramondi Bryopsis hypnoides, Centroceras clavulatum, Ceramium flaccidum, Derbesia vaucheriaeformis, Enteromorpha prolifera (Ulva), Giffordia rallsiae, Polysiphonia havanensis --- Washington, D.C
Appendices D-5 

Table D.2. Summary of the most frequently found macroalgae divisions and functional form groups that 

were found in foraging samples of juvenile green turtles (Chelonia mydas) and herbivorous 

fishes. Turtle data are derived from studies listed in Appendix A, Table A.1, and fish data 

are from footnoted sources on Table D.3. 

Number of 

Herbivorous 

Fish/Turtles 

Macroalgal Species Functional Form Group Division 

9 Ceramium sp. Filamentous Rhodophyta 

9 Ulva sp. Sheet Chlorophyta 

8 Polysiphonia sp. Filamentous Rhodophyta 

8 Lyngbya sp. Filamentous Cyanobacteria 

7 Centroceras sp. Filamentous Rhodophyta 

7 Dictyota sp. Sheet Phaeophyta 

6 Amphiroa sp. Jointed-calcareous Rhodophyta 

6 Coelothrix sp. Coarsely-branched (fish only) Rhodophyta 

6 Herposiphonia sp. Coarsely-branched Rhodophyta 

6 Hypnea sp. Filamentous Rhodophyta 

6 Laurencia sp. Coarsely-branched Rhodophyta 

5 Gelidium sp. Coarsely-branched Rhodophyta 

5 Caulerpa sp. Coarsely-branched Chlorophyta 

5 Jania sp. Jointed-calcareous Rhodophyta 

Table D.3. Macroalgal species in foraging samples from different population studies conducted on 

juvenile green turtles (Chelonia mydas) and herbivorous fish populations on nearshore 

hardbottom habitat. Data are derived from Appendix A, Table A.1 and footnoted sources. 

Macroalgal Species 

Chelonia mydas Acanthurus Scartella Entomacrodus Ophioblennius Kyphosus Holocanthus Pomocanthus Abudefduf Microspathodon Stegastes Sparisoma Archosargus Diplodus Lagodon 

Acanthophora muscoides √ √ 

Acanthophora sp. √ √ 

Acanthophora spicifera √ 

Agardhiella subulata √

Amphiroa fragillisma √ 

Amphiroa rigida √ 

Amphiroa sp. √ √ √ √ 

Asteromenia (Fauchea peltata) √ 

Avrainvillea sp. √ 

Bostrichia sp. √ 

Botryocladia occidentalis √

Botryocladia sp. √ √ 

Bryocladia cuspidata √ 

Bryopsis pennata √ 

Bryopsis sp. √ √ √ 

Bryothamnion seaforthii √ 

Bryothamnion sp. √ √ 

Callithamnion sp. √ 

Caulerpa cupressoides √ 

Caulerpa mexicana √ 

Caulerpa microphysa √

Caulerpa peltata √
Table D.3. (Continued). 

Appendices D-6 

Macroalgal Species 

Chelonia mydas Acanthurus Scartella Entomacrodus Ophioblennius Kyphosus Holocanthus Pomocanthus Abudefduf Microspathodon Stegastes Sparisoma Archosargus Diplodus Lagodon 

Caulerpa prolifera 

√ 

Caulerpa racemosa 

√ √ 

Caulerpa sertularioides 

√ 

Caulerpa sp.

√√ √√ 

Caulerpa taxifolia 

√ √ 

Centroceras clavulatum 

√ √√ 

Centroceras sp.

√ √√√ 

Ceramium byssoideum 

√ 

Ceramium sp. 

√√√ √ √√√√ 

Chaetomorpha linum 

√ 

Chaetomorpha sp.

√ √ 

Champia parvula 

√ 

Champia sp. 

√ 

Chondria dasyphylla 

√ 

Chondria sp.

√ √ √ 

Chondrocanthus acicularis 

√ 

Cladophora catenata 

√

Cladophora delicatula 

√ 

Cladophora sp.

√√ √ 

Codium isthmocladum 

√ 

Codium sp.

√ 

Coelothrix irregularis 

√√ 

Coelothrix sp. 

√ √√ √ 

Colpomenia sp. 

√ 

Cryptonemia crenulata 

√

Dascycladus vermicularis 

√ 

Dasya pedicillata 

√

Dasya sp.

√ 

Dictyopteris delicatula 

√ √√ 

Dictyopteris plagiogramma 

√ 

Dictyopteris sp.

√ √√ 

Dictyospareia sp.

√√ 

Dictyota bartayresii 

√ √ 

Dictyota dentata 

√ 

Dictyota divaricata 

√ 

Dictyota sp.

√√√ √√ 

Digenia sp. 

√ 

Dudresyna crassa 

√ 

Ectocarpus sp.

√ 

Enteromorpha sp.

√ 

Erythrotrichia carnea 

√ 

Eucheuma nudum 

√

Eucheuma sp. 

√ √ 

Galaxaura sp. 

√ 

Gelidiaceae 

√ 

Gelidiella sp.

√ √ 

Gelidium americanum 

√ 
Table D.3. (Continued). 

Appendices D-7 

Macroalgal Species 

Chelonia mydas Acanthurus Scartella Entomacrodus Ophioblennius Kyphosus Holocanthus Pomocanthus Abudefduf Microspathodon Stegastes Sparisoma Archosargus Diplodus Lagodon 

Gelidium pusillum 

√ 

Gelidium rigidulum 

√ 

Gelidium sp.

√√ √ √ 

Gracilaria armata 

√ 

Gracilaria blodgettii 

√

Gracilaria mammillaris 

√

Gracilaria sp.

√ √ 

Gracilaria tikvahiae 

√ 

Gracilaria verrucosa 

√

Halimeda discoidea 

√

Halimeda sp. 

√√ 

Halymenia florisia 

√ √ 

Herposiphonia sp.

√√√ √√√ 

Hypnea cervicornis 

√ 

Hypnea cornuta 

√

Hypnea musciformis 

√ √ 

Hypnea sp.

√√√ √√ 

Hypnea spinella 

√ √ 

Hypnea valentiae 

√ 

Jania adherens 

√ √ 

Jania rubens 

√ 

Jania sp.

√√ √ √ 

Laurencia corallopsis 

√ 

Laurencia gemmifera 

√

Laurencia obtusa 

√ 

Laurencia papillosa 

√ 

Laurencia poiteaui 

√ 

Laurencia sp. 

√√√ √ 

Liagora sp.

√ 

Lomentaria baileyana 

√ 

Neomeris annulata 

√

Nitophyllum punctatum 

√

Padina gymnospora 

√ 

Padina profunda 

√ 

Padina sp.

√ √ 

Penicillus sp. 

√ 

Pocockiella sp.

√ √√ 

Polysiphonia sp.

√√√√ √√√√ 

Polysiphonia subtilissima 

√ 

Pterocladia sp.

√ 

Rhizoclonium sp.

√√ √√ 

Rhodymenia sp.

√ 

Sargassum platycarpum 

√ 

Sargassum sp.

√√ √ 

Scinaia complanata 

√ 

Scinaia sp.

√ 

Siphonocladus tropicus 

√ 
Table D.3. (Continued). 

Appendices D-8 

Macroalgal Species 

Chelonia mydas Acanthurus Scartella Entomacrodus Ophioblennius Kyphosus Holocanthus Pomocanthus Abudefduf Microspathodon Stegastes Sparisoma Archosargus Diplodus Lagodon 

Solieria sp. √ 

Spatoglossum sp. √ 

Sphacelaria sp. √ 

Spyridia filamentosa √ √ 

Spyridia sp. √ √ 

Turbinaria sp. √ 

Ulva lactuca √ 

Ulva sp. √ √ √ √ √ √ √ √ 

Ulva sp. (Enteromorpha chaetomorphoides) √ 

Wrangelia argus √ 

Cyanobacteria 

Anacystis sp. √ √ √ 

Calothrix sp. √ 

Lyngbya majuscula √ 

Lyngbya sp. √ √ √ √ √ √ √ 

Microcoleus sp. √ √ √ √ 

Oscillatoria sp. √ √ 

Phormidium sp. √ √ 

Plectonema nostocorum √ 

Plectonema sp. √ 

Schizothrix arenaria √ 

Schizothrix calcicola √

General Categories 

Plants and algae √1 

Algae √2 √2, 3 √7 √8 √7, 9

Benthic algae/weeds √4 

Detritus & diatoms √5 

Filamentous green √6 

1 Dominici-Arosemena and Wolff, 2005.

2 Hourigan et al., 1989. 

3 Feddern, 1968. 

4

 Randall, 1967. 

5 Randall, 1965. 

6 Irvine, 1982 in Gerking ,1994. 

7

 Hay and Sutherland, 1988. 

8 Carr and Adams, 1972; Pike and Lindquist, 1994; Hay and Sutherland, 1988. 9 Motta et al., 1995.