OCS Study 

MMS 95-0029 

Synthesis of Available Biological, 

Geological, Chemical, Socioeconomic, 

and Cultural Resource Information 

for the South Florida Area 

Supplemental Report: A Comparison of 

Seagrass Beds in Panama and South Florida 

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U .S . Department of the Interior 

Minerals Management Service 

Gulf of Mexico OCS Region 
OCS Study 

MMS 95-0029 

Synthesis of Available Biological, 

Geological, Chemical, Socioeconomic, 

and Cultural Resource Information 

for the South Florida Area 

Supplemental Report : A Comparison of 

Seagrass Beds in Panama and South Florida 

Prepared by 

Continental Shelf Associates, Inc. 

Prepared under MMS Contract 

14-35-0001-30417 

by 

Continental Shelf Associates, Inc. 

759 Parkway Street 

Jupiter, Florida 33477 

Published by 

U .S . Department of the Interior 

Minerals 1management Service New Orleans 

Gulf of Mexico OCS Region May 1995 
DISCLAIMER 

This report was prepared under contract between the Minerals Management 

Service (MMS) and Continental Shelf Associates, Inc. This report has been 

technically reviewed by the MMS and approved for publication. Approval does 

not signify that the contents necessarily reflect the views and policies of the 

Service, nor does the mention of trade names or commercial products constitute 

endorsement or recommendation for use. It is however, exempt from review and 

compliance with MMS editorial standards. 

REPORT AVAILABILITY 

Extra copies of this report may be obtained from the Public Information Unit 

(Mail Stop 5034) at the following address: 

U.S . Department of the Interior 

Minerals Management Service 

Gulf of Mexico OCS Region 

Public Information Unit (MS 5034) 

1201 Elmwood Park Boulevard 

New Orleans, Louisiana 70123-2394 

Telephone Number: 1-800-200-GULF or 

(504) 736-2519 

CITATION 

Continental Shelf Associates, Inc. 1995. Synthesis of Available Biological, 

Geological, Chemical, Socioeconomic, and Cultural Resource Information 

for the South Florida Area. Supplemental Report: A Comparison of 

Seagrass Beds in Panama and South Florida. OCS Study MMS 95-0029. 

U.S . Dept. of the Interior, Minerals Mgmt. Service, Gulf of Mexico OCS 

Region, New Orleans, LA. 33 pp. + app. 

iii 
TABLE OF CONTENTS 

Page 

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

LIST OF TABLES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .vii 

ACKNOWLEDGMENTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ix 

INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .1 

Background: The Bahia Las Minas Spill . . . . . . . . . . . . . . . . . . . . . 1 

Subtidal Seagrass Study . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 

Other Study Components . . . . . . . . . . . . . . . . . . . . . . . . . . 6 

Study Approach . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 

SEAGRASS DISTRIBUTION AND ENVIRONMENTAL SETTING . . . . . . . . . 9 

Geographic Distribution and Areal Extent . . . . . . . . . . . . . . . . . . . 9 

Water Depth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 

Proximity to Mangrove Shorelines . . . . . . . . . . . . . . . . . . . . . . . . 11 

Wave Energy Regime . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 

History of Environmental Problems . . . . . . . . . . . . . . . . . . . . . . . 13 

SPECIES COMPOSITION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 

Flora . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 

Seagrasses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 

Algae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 

Fauna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 

Decapods and Stomatopods . . . . . . . . . . . . . . . . . . . . . . . 17 

Molluscs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 

Fishes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 

Other Fauna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 

CONCLUSIONS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .25 

Seagrass Distribution and Environmental Setting . . . . . . . . . . . . . 25 

Species Composition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 

Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 

REFERENCES CITED . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 

APPENDIX: SPECIES LISTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A-1 

v 
LIST OF FIGURES 

Figure Description Page 

1 Study area . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 

2 Seagrass study sites in the region of Panama affected 

by the April 1986 oil spill (From: Marshall et al . 

1993) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3 

LIST OF TABLES 

Table Description Page 

1 Summary of faunal comparisons between seagrass 

beds of South Florida and Panama . . . . . . . . . . . . . . . . . . . . . 18 

2 Numerically abundant caridean and penaeid shrimp 

from seagrass beds of South Florida (Florida 

Bay--Holmquist et al . 1989) and Panama (Bahia Las 

Minas area--M. Marshall unpubl . data) . . . . . . . . . . . . . . . . . . 19 

3 Numerically abundant fish species from seagrass 

beds of South Florida (Cape Romano area) and 

Panama (Bahia Las Minas area) . . . . . . . . . . . . . . . . . . . . . . . 22 

A.1 Comparison of decapod and stomatopod crustacean 

species lists for South Florida (Florida Bay) and 

Panama (Bahia Las Minas area) . . . . . . . . . . . . . . . . . . . . . . A-3 

A.2 Comparison of mollusc species lists for South 

Florida (mainly Florida Bay) and Panama (Bahia 

Las Minas area) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A-5 

A.3 Comparison of fish species lists for South Florida 

and Panama . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A-11 

vii 
ACKNOWLEDGMENTS 

This Supplemental Report was prepared by Dr. Neat W. Phillips (CSA) from 

an original draft written by Dr. Michael J. Marshall (now with Mote Marine 

Laboratory) and Mr. M. John Thompson (CSA). Dr. Richard M. Hammer 

(CSA) was the Program Manager. Mr. David B. Snyder (CSA) reviewed the fish 

section and provided additional data for the Panama-South Florida comparison . 

The copy editor was Ms. Melody Powell (CSA) . 

Dr. Joseph C. Zieman (University of Virginia) and Dr. Marshall reviewed an 

earlier draft of the Supplemental Report and provided helpful, constructive 

comments. Report conclusions, as well as any remaining errors or technical 

inaccuracies, are the responsibility of Continental Shelf Associates, Inc. 

ix 
INTRODUCTION 

Numerous environmental and socioeconomic studies have been conducted in 

South Florida by government agencies, university researchers, private organizations, and individuals. In 1988, the Minerals Management Service (MMS) 

awarded a contract to Continental Shelf Associates, Inc. to synthesize the available information and evaluate potential effects of offshore oil and gas exploration 

and development. The goal was to help policy makers reach informed decisions 

about future lease offerings and environmental restrictions on offshore oil and 

gas operations . In May 1990, a Final Report entitled "Synthesis of Available 

Biological, Geological, Chemical, Socioeconomic, and Cultural Resource 

Information for the South Florida Area" (Phillips and Larson 1990) was 

completed. A Master Bibliography and an Executive Summary were 

subsequently produced. The study area is shown in Figure 1. 

Oil spill effects on South Florida seagrass, coral reef, and mangrove 

communities were a key topic in the Final Report. Because there have been no 

major oil spill studies in South Florida, results from similar environments 

elsewhere were reviewed. One of the most frequently cited examples was the 

1986 Bahia Las Minas spill in Panama, which was the largest ever recorded in 

coastal habitats of the tropical Americas (Jackson et al. 1989). The spill occurred 

on the Atlantic coast near the field laboratory of the Smithsonian Tropical 

Research Institute (STRI). Post-spill studies by STRI scientists began 

immediately, with funding by the MMS. A synthesis report was issued in 1993 

(Keller and Jackson 1993). 

The MMS was interested in the Bahia Las Minas spill because study findings 

might help to predict oil spill impacts in South Florida. The coastal 

environments and many of the species affected in Panama are similar to those 

found in South Florida. Jackson et al . (1989), reporting preliminary observations 

of the Panama spill, noted that "our observations are relevant to assessment of 

potential biological effects of pollution in several areas where extraction or 

refining of oil is ongoing or planned," including South Florida. However, no one 

has systematically compared conditions in Panama and South Florida to 

determine whether and to what extent study results are transferable . That critical 

question is the topic of this Supplemental Report. This comparison focuses on 

seagrass communities. 

BACKGROUND: THE BAHIA LAS MINAS SPILL 

In April 1986, 9.6 to 16.0 million liters (60,000 to 100,000 barrels) of 

medium-weight crude oil spilled from a refinery storage tank into Panamanian 

coastal waters (Keller and Jackson 1993). The spill occurred in a complex region 

of seagrass beds, coral reefs, and mangrove forests near the Caribbean entrance 

to the Panama Canal (Figure 2). For six days after the spill, onshore winds 

caused the oil to remain within the small embayment (Bahia Cativa) adjacent to 

the refinery. Then, shifting winds and runoff from rains caused a large quantity 

of oil to float to sea past a boom across the mouth of the embayment (Keller and 
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spill (From: Marshall et al. 1993) . A. General location in Panama. B. Location of 

unoiled sites (BNV, DONT, PALN, and LINE). The boxed area indicates the site of the 

1986 spill. C. Location of the refinery (R) and oiled sites in Bahia Las Minas (MINK, 

PGN, LREN, and LRS). 

3 
Introduction 

Jackson 1993). Subsequently, the oil spread along the coast, reaching mangroves, 

small estuaries, and sand beaches within 10 km of the refinery. 

The spill site was near the Galeta Marine Laboratory of the STRI, and some 

of the oiled environments had been studied previously by STRI scientists . The 

MMS and the STRI quickly entered into a cooperative agreement to investigate 

the immediate and long-term effects of this spill. A synthesis report was issued 

in 1993 (Kelley and Jackson 1993). The following discussion of STRI seagrass 

studies is based on the synthesis report (Kelley and Jackson 1993), especially the 

chapter on subtidal seagrass communities (Marshall et al. 1993). 

Subtidal Seagrass Study 

The Panamanian seagrass beds oiled by the 1986 spill occur within a 11 km 

straight line distance extending from Isla Margarita to the west to las Islas 

Naranjos on the eastern edge of Bahia Las Minas (Figure 2) . The actual shoreline distance affected by the spill, due to the geographical complexity created by 

a labyrinthine system of small islands, river mouths, and numerous embayments, 

is about 82 km (Jackson et al. 1989). The seagrass beds were 1 to 2 ha in size 

and were located in shallow lagoons (<1 m depth) between small fringing reefs 

and mangrove-covered shorelines. 

Seagrass bed studies completed before the spill include epifaunal (Heck 1977, 

1979; Weinstein and Heck 1979) and infaunal surveys (Jackson 1973; VasquezMontoya 1979, 1983). However, the pre-spill data were not appropriate as a 

quantitative baseline for evaluating effects (Marshall et al. 1993). Therefore, 

effects on seagrasses were evaluated by comparing oiled and unoiled sites after 

the spill. Four oiled and four unoiled sites (Figure 2) were sampled quarterly 

from September 1986 until July 1989. All oiled sites were within 6 km of the 

refinery. The unoiled sites were located a considerable distance from the 

refinery (20 to 40 km), but were believed to be similar to the oiled beds in 

physical and biological characteristics (Marshall et al. 1993) . 

Water depth profiles were produced initially for each site along transects 

from the shore to the seaward edge of each bed. The investigators also 

measured the position of the seaward edge of seagrass beds after July 1988 (when 

they first noticed that some oiled beds appeared to be receding) . Background 

data collected for each site included salinity, sediment silt/clay percentage and 

organic content, and presence/absence of visible oil. On each sampling date, core 

samples of seagrass, macroalgae, and benthic infauna were collected, and surface 

sediment scrapes were obtained for grain size and hydrocarbons. Mobile 

epifauna were also collected at each site, using pushnet samplers. Infauna and 

epifauna were identified to major taxonomic groups (except for polychaetes, 

which were identified to family, and caridean shrimp, which were identified to 

species) . Size frequency and reproductive characteristics were measured on the 

two most abundant species of caridean shrimp. 

4 
Introduction 

Marshall et al . (1993) reported findings of the subtidal seagrass study. Their 

major points are summarized below: 

Oil was deposited on and persisted in seagrass bed sediments. 

Concentrations as high as 4,020 ppm were detected in surface sediments 

five months after the spill. Two years later, analyses showed that oil had 

penetrated thick mats of seagrass roots and rhizomes to a depth of at 

least 20 cm (Burns 1993). 

Seagrasses died at the shoreward margins of some oiled seagrass beds. At 

one site visited two years after the spill, seagrass had died within a 5-m 

wide band along the shoreward margin. It is unknown whether similar 

losses occurred at other sites prior to this observation. Subsequently, 

changes in seagrass bed margins were measured at all sites for about one 

year, and the margins of oiled beds generally receded (20 to 90 cm), 

whereas unoiled beds generally did not. 

The spill apparently had a transitory effect on the predominant seagrass, 

Thalassia (other than at the bed margins, as noted above). Total, 

subsurface, and blade biomass of Thalassic was lower initially at oiled 

sites. However, this difference disappeared within a few months, and 

subsequently, seagrass biomass at both oiled and unoiled sites generally 

decreased through the last sampling period, approximately three years 

after the spill . Syringodium, the second-ranked seagrass, was more 

sensitive than Thalassic to oiling; Syringodium biomass remained lower at 

oiled sites even during the second and third years after the spill. 

Living calcareous algae were virtually absent at oiled sites initially, but 

this difference did not persist. The abundance of fleshy algae did not 

seem to be affected by oiling . 

Total densities of infauna (excluding polychaetes) and sedentary epifauna 

were initially lower in oiled beds than in unoiled beds. During the three 

years after the spill, abundance generally increased at oiled sites and 

decreased at unoiled sites (the latter for unknown reasons), eventually 

converging to similar levels . Amphipods, cumaceans, isopods, ophiuroids, 

sipunculids, and tanaids appeared to be strongly affected, whereas 

gastropods, bivalves, and brachyuran crabs apparently were not. 

Among epifauna, groups with direct development were more strongly 

affected than were partial brooders or those with pelagic larvae. This 

might have been due to effects of oiling on reproduction of brooding 

species within the bed. In contrast, species with pelagic larvae could have 

repopulated through recruitment from distant seagrass beds. 
I ntroduction 

Epifaunal echinoderms were strongly affected by the spill; large sea 

urchins and holothurians were still more abundant at unoiled sites six 

years after the spill. Because these organisms are easily counted in the 

field, the investigators suggested that they may be useful as "indicators" of 

the effects of oil spills in seagrass beds. 

Other Study Components 

In addition to the subtidal seagrass study, several other components of the 

STRI project provide relevant information to evaluate the applicability of the 

Panama spill data to South Florida: 

Hydrocarbon data show that oil persisted in mangrove forest sediments 

for more than five years (Burns 1993) . Although much of the spilled oil 

weathered rapidly, oil buried at one mangrove site was preserved 

relatively intact in anoxic sediments for five years. Oil in mangrove 

sediments constitutes a source of chronic, potentially toxic leakage onto 

adjacent habitats, including seagrass beds. 

One of the most dramatic effects of the spill was the destruction of an 

intertidal seagrass bed/reef flat habitat at one site (Steger and Caldwell 

1993). Before the spill, there was an intertidal seagrass bed with scattered 

pieces of coral rubble, bordered by a mangrove stand. After the seagrass 

was killed by oil and decayed, the coral rubble and sediment formerly 

bound together by roots and rhizomes began to erode. The area is now 

dominated by large pieces of coral rubble resting on a fine, sand/silt 

substratum, bordered by a mud flat . There is little chance that this 

habitat will ever recover. 

About 64 ha, or 7% of the mangrove habitat in the area, was deforested, 

mostly in a 50-m wide coastal strip (Duke and Pinzon 1993). A much 

larger area was affected to some extent by oiling. Mature trees surviving 

the initial oiling suffered persistent canopy deterioration . In oiled canopy 

gaps, seedling recruitment and growth were extremely variable, with 

significant growth suppression in areas of residual oil . These effects 

persisted at least 5 to 6 years after the spill . Loss of mangrove forest 

resulted in erosion and increased sedimentation on adjacent seagrass beds 

and coral reefs. 

There was a striking reduction in cover, abundance, and diversity of live 

corals (especially the elkhorn coral Acropora palmata) immediately after 

the spill (Guzman et al . 1993) . On oiled reefs, coral injuries were more 

frequent and coral growth rates were lower than on unoiled reefs. Low 

coral recruitment rates on oiled reefs indicated there was little prospect 

for rapid recovery . 

6 
Introduction 

STUDY APPROACH 

The comparison of seagrass beds in South Florida and Panama is not 

intended to be exhaustive. Rather, it focuses on those aspects that are most 

critical in predicting the effects of oil spills on seagrasses. The response of 

seagrass beds to oiling depends on a variety of environmental factors (Zieman et 

al. 1984; Marshall et al . 1990) . For this report, the factors are divided into three 

groups : 

Swill Circumstances. The effects of an oil spill depend in part on 

unpredictable factors such as spill location, volume, duration, rate of 

release, and chemical composition, as well as the effectiveness of spill 

containment and cleanup measures. Effects also depend on unpredictable 

environmental variables such as weather, sea state, and tidal stage at the 

time of a spill. 

Seaerass Distribution and Environmental Setting. Factors that 

determine the severity of spill effects include the geographic distribution, 

areal extent, and water depth of seagrass beds; bed location, including 

proximity to mangrove shorelines; tidal range and wave energy regime; 

and history of environmental problems (Zieman et al. 1984; Jackson et al. 

1989; Marshall et al. 1990). 

Species Composition of seagrass bed communities, including both seagrasses and associated flora and fauna. Species composition is important 

because species differ in their sensitivity to oiling (Gilfillan 1990). 

This report focuses on the last two topics, which are discussed in separate 

chapters . Spill circumstances are not part of the comparison because they cannot 

be predicted in advance. The exact circumstances of the Bahia Las Minas spill 

could not be repeated in the South Florida study area because no oil refineries 

are present, nor are they likely to be permitted in the coastal zone (Phillips 

1990). However, there is a large volume of tanker traffic passing through the 

Straits of Florida, which could produce a spill of comparable or greater volume. 

7 
SEAGRASS DISTRIBUTION AND 

ENVIRONMENTAL SETTING 

GEOGRAPHIC DISTRIBUTION AND AREAL EXTENT 

South Florida possesses one of the largest seagrass resources in the world; 

total coverage of seagrass beds in South Florida is estimated to be about 

5,500 km2 (Zieman 1982; Iverson and Bittaker 1986; Zieman and Zieman 1989). 

The South Florida seagrass beds range in size from thousands of hectares to 

small patches in mangrove-lined bays and lagoons . The seagrasses occur in four 

main areas: (1) throughout Florida Bay', which provides a shallow, protected 

environment for extensive seagrass development; (2) Hawk Channel between the 

Florida Keys and the Florida Reef Tract; (3) the lower Keys from Big Pine Key 

to Key West; and (4) the Marquesas-Dry Tortugas area. Greater water clarity on 

the Atlantic side of the Florida Keys allows extensive seagrass bed development 

in Hawk Channel, despite the higher wave energy. To the north of the Florida 

Keys on the Atlantic coast, increasing wave exposure limits seagrass beds to small 

pockets in protected inlets and lagoons (Zieman 1982). Turbidity and reduced 

salinity due to Everglades drainage limit seagrass bed distribution along the Gulf 

Coast north of Florida Bay (Zieman and Zieman 1989), although sparse 

Halophila beds occur offshore on the Southwest Florida shelf (Continental Shelf 

Associates, Inc. 1989). 

In contrast, the steep terrain of the nearshore shelf in Panama prevents the 

development of extensive offshore seagrass beds. Additionally, the narrow 

continental shelf of Panama's Caribbean coast renders this area a higher energy 

coastline than most of South Florida. Seagrass beds are generally located in 

protected waters, such as in island-fringed lagoons or estuaries, or behind reefs 

or barrier islands (Zieman 1982, 1990). Consequently, Panamanian seagrass beds 

occur mainly behind well developed reefs and barrier islands where these structures absorb wave energy and allow seagrass development . Panamanian seagrass 

beds occur in small, discontinuous patches often separated by wide areas of coral 

reef or sandy channel (Heck 1979; M. Marshall, pers. observ.) . 

None of the Panamanian seagrass beds studied were larger than 2 ha in size, 

because there are no larger beds along this coast. These beds are not 

comparable to the much larger seagrass meadows typical of Florida Bay and the 

Florida Keys. South Florida seagrass beds can extend for many kilometers 

without being interrupted by other habitats such as mud banks, oyster bars, coral 

reefs, or tidal channels. 

This comparison shows that South Florida seagrass beds are extensive, 

whereas the Panamanian beds are small and discontinuous. The difference is 

1 Florida Bay as discussed here corresponds to the boundaries of Everglades National Park south 

of Cape Sable. 

9 
Seagrass Distribution and Environmental Setting 

important from at least two respects. First, despite the size of the Panama spill 

and the length of coastline affected, a relatively small area of seagrass beds was 

contaminated. A much larger area of South Florida seagrass beds could be oiled 

by a single spill. Second, the pattern of recovery from an oil spill might be 

different in South Florida, because some animals could migrate to unaffected 

areas and extensive source beds for new recruits would exist. Small, isolated 

seagrass beds may be subject to severe setbacks or total elimination, as 

contrasted with seagrass beds that are part of a larger, continuous system 

(Zieman et al . 1984). However, incorporation of oil in seagrass bed sediments 

(as happened in Panama) could inhibit recovery even if new recruits were 

available. 

WATER DEPTH 

Seagrass vertical distribution is limited by exposure and desiccation in shallow 

water and by the availability of photosynthetically active radiation at greater 

depths. In South Florida, seagrasses grow at depths ranging from the intertidal 

to >40 m (Zieman 1982; Continental Shelf Associates, Inc. 1989) . However, the 

development of rich, productive seagrass beds is generally limited to water depths 

of < 10 to 12 m (Zieman 1982, 1990). In Florida Bay, which is characterized by a 

"honeycomb" pattern of anastomosing mudbanks (shoals) and basins, maximum 

seagrass density occurs at depths of 1 to 3 m. Extensive seagrass beds also occur 

in water depths of 1 to 3 m around islands in the lower Keys. In the clearer 

waters of Hawk Channel behind the Florida Reef Tract, dense beds occur at 

depths of 6 to 8 m (Zieman 1990). 

All of the Panamanian seagrass beds studied by Marshall et al . (1993) are in 

shallow water (< 1 m; mean depth ranged from 11 to 64 cm) and are situated 

between mangrove shorelines and fringing coral reefs. The steep terrain of the 

nearshore shelf in Panama prevents the development of extensive offshore 

seagrass beds in deeper water. Seagrasses also occur on intertidal reef flats in 

Panama (Jackson et al . 1989; Steger and Caldwe111993). 

With respect to depth distribution of seagrasses, the Panamanian sites studied 

by Marshall et al. (1993) seem most similar to shallow, subtidal seagrass beds in 

Florida Bay and adjacent to islands in the lower Florida Keys (M. Marshall, pens. 

observ. ; M. J. Thompson, pegs. observ. ; J. Zieman, 1992, pets. comm., Univ. of 

Virginia) . 

According to Marshall et al . (1990), intertidal and shallow subtidal seagrass 

beds are the most susceptible to oiling. The most severe effects on seagrasses in 

the Panama spill study occurred on intertidal reef flats (Jackson et al . 1989; 

Steger and Caldwell 1993). Heavily oiled intertidal beds were killed and may 

never recover (Steger and Caldwell 1993), whereas subtidal seagrass beds 

survived everywhere, with mortality at the margins and more subtle effects within 

the beds (Marshall et al . 1993). 

10 
Seagrass Distribution and Environmental Setting 

Data from subtidal Panamanian seagrass beds (Marshall et al . 1993) could 

greatly underestimate the severity of oil spill effects on Florida Bay seagrasses. 

Many seagrasses on Florida Bay mudbanks are exposed at very low tides and 

could be highly vulnerable to oiling, similar to the intertidal reef flat beds in 

Panama studied by Steger and Caldwell (1993) . The mudbanks are concentrated 

in the western part of Florida Bay, and the shallowest portions of these beds 

support a seagrass standing crop that is twice the bay-wide average (Zieman 

1990). 

On the other hand, data from the shallow subtidal Panamanian beds may 

overestimate oil spill impacts on Hawk Channel seagrasses. Because of their 

greater water depth, the Hawk Channel beds might escape oil exposure. 

However, under certain conditions (e.g., storms, wave action), oil can become 

mixed into emulsions with seawater and combine with sediment particles and sink 

to the bottom, resulting in oiling of deeper beds (Marshall et al . 1990). 

PROXIMITY TO MANGROVE SHORELINES 

All of the Panamanian seagrass beds studied occur in shallow lagoons 

between fringing coral reefs and mangrove shorelines. The distance between 

seagrass beds and mangrove shoreline ranged from a few meters to a few tens of 

meters (Marshall et al. 1993). This is significant because mangrove sediments 

tend to absorb spilled oil, which may then be released gradually over a period of 

years. Hydrocarbon data show that oil persisted in mangrove forest sediments 

for more than five years after the spill (Burns 1993). Although much of the 

spilled oil weathered rapidly, oil buried at one mangrove site was preserved 

relatively intact in anoxic sediments for five years. Oil slicks, presumably from 

mangrove sediments, were seen in Bahia Las Minas throughout the duration of 

the seagrass project. This oil constitutes a source of chronic, potentially toxic 

leakage onto adjacent seagrass beds. 

From the standpoint of mangrove shorelines, the Panamanian seagrass beds 

are more similar to those of Florida Bay and the lower Keys rather than those of 

Hawk Channel. Extensive mangrove forests occur on the mainland coast of 

Florida Bay, and mangroves also occur on islands throughout the bay and on the 

shoreline of the Florida Keys. Some seagrass beds in Florida Bay and the lower 

Keys are near mangrove shoreline, whereas the Hawk Channel beds are not. 

Both the seagrass beds and the mangrove forests of South Florida are much 

larger and more extensive than those of Panama (Snedaker 1990; Zieman 1990). 

Although most individual seagrass beds would not be as close to mangrove 

shorelines as those in Panama, a similar effect could be observed if a major spill 

reached Florida Bay or the lower Keys. Oil reaching the mangrove shorelines 

would be absorbed and would probably leak out slowly over many years, as 

happened in Panama. 

11 
Seagrass Distribution and Environmental Setting 

WAVE ENERGY REGIME 

The importance of wave exposure in limiting seagrass bed development has 

been cited above. Wave energy is also important from other respects in 

determining the effects of an oil spill. For example, wave action can help to 

break up oil slicks, and it can enhance the effectiveness of dispersants2 . Two 

other important aspects are discussed below. 

Wave action can enhance sedimentation of oil by creating oil-in-water 

emulsions that settle with sediment particles . In August 1986, several months 

after the Bahia Las Minas spill, an emulsion of oil suspended in seawater was 

seen over the fringing reefs at the seaward edges of two seagrass study sites 

(Marshall et al. 1993). The emulsion presumably was created by turbulence 

resulting from waves crashing onto the reef crest. Droplets from this emulsion 

may have combined with suspended sediment particles to produce a 

heavier-than-water combination of oil and sediment. Oil sunken by this 

mechanism may have accounted for some part of that detected in seagrass bed 

sediments. Because the Panama seagrass beds are on a high energy coast, 

compared to the usually calm waters of the vast expanses of Florida Bay, this 

proposed mechanism may not be as important over the South Florida seagrass 

beds. However, winter storms in South Florida are energetic and could produce 

a similar effect in Florida Bay. 

Wave regime can also influence the resilience of seagrass communities 

following a disturbance. It has been noted above that the Panamanian seagrass 

beds occur along a higher energy coast than South Florida. Perhaps for this 

reason, Panamanian seagrass beds seem more susceptible to wave induced 

erosion (blowouts) than do the large seagrass beds off Florida (Heck 1979). 

Such blowouts occur when normal, dry season wave action breaches the integrity 

of the root system within a seagrass bed and produces an open, sandy area. 

Once opened, the seaward edges of such blowouts usually continues to erode 

toward the sea, while the shoreward edge tends to repopulate with seagrass 

(Patriquin 1975). The results of this phenomenon render the seagrass habitats 

off Panama more dynamic in terms of species succession and regeneration than 

their counterparts off South Florida. 

Z About one week after the Bahia Las Minas spill, an aircraft sprayed approximately 21,000 liters 

of the dispeisant Corexit 9527 onto oil slicks . According to Keller and Jackson (1993), "the 

application of dispersant so many days after the spill and the calm sea conditions during the spraying 

appeared to render chemical dispersion ineffective." 

12 
Seagrass Distribution and Environmental Setting 

HISTORY OF ENVIRONMENTAL PROBLEMS 

Both the Panamanian and South Florida coasts have been affected by human 

disturbance over a period of many years. These disturbances may have altered 

the sensitivity and resilience of seagrass beds to oiling . As noted by Jackson et 

al . (1989), the response of organisms to an oil spill depends in part on the 

environmental conditions (including chronic disturbance and pollution) under 

which they exist. 

Much of Bahia Las Minas has been subjected to human disturbance, 

including decades of excavation, dredging and filling for construction of the 

Panama Canal and the city of Colon; construction of a refinery, a cement plant, 

and an electrical generating station; drainage and spraying of mangroves for 

mosquito control; extensive deforestation, resulting in deposition of eroded 

terrigenous sediments in coastal environments ; a previous major oil spill in 1968 

(the wreck of the tanker Witwater) ; and numerous minor spills from vessels and 

port facilities (Jackson et al . 1989; Keller and Jackson 1993). Effects of the 1968 

Witwater spill on mangrove forests were still noticeable 20 years later (Duke and 

Pinzon 1993). The cumulative effects of these disturbances (if any) on seagrass 

communities are unknown. However, the eventual convergence of seagrass bed 

faunal populations at oiled and unoiled seagrass sites in the years after the 1986 

spill suggests that pre-spill status was similar throughout the area (Marshall et al. 

1993). 

Coastal and nearshore environments of South Florida have also faced 

numerous anthropogenic problems that have led to decreased seagrass acreage. 

Historically, much seagrass habitat in South Florida has been lost to dredging 

and filling for residential and commercial development (Zieman 1982). 

According to Zieman (1990), the main cause for continued seagrass losses in 

South Florida is increased turbidity, attributable either to sedimentation (e.g., 

from construction activities) or eutrophication (e.g ., from inadequate sewage 

treatment in the Florida Keys). Other problems include physical damage 

(scarring) by small boats and chronic pollutant releases from vessels of all types. 

Severe water quality and ecological problems have developed in Florida Bay 

in recent years. Problems include a massive seagrass die-off, phytoplankton 

blooms, sponge die-offs, mangrove die-backs, and population reductions in 

economically significant species such as pink shrimp, sponges, lobster, and various 

recreational gamefish. Although the causes and mechanisms are not fully 

understood (Boesch et al . 1993), changes in freshwater inflow from the 

Everglades are believed to be a major influence (McIvor et al . 1994). 

The history of environmental degradation may affect the response of seagrass 

communities to oiling. In some situations, chronic exposure to pollutants and 

other environmental stresses produces altered communities that are resistant to 

disturbance. However, it is not clear what role, if any, the previous 

environmental stresses played in determining the effects of the Panama spill. 

13 
SPECIES COMPOSITION 

FLORA 

Seagrasses 

The species composition of seagass communities depends on the seagrasses 

present and their respective densities. Thalassic testudinum (turtle grass), 

Syringodium fcliforme (manatee grass), and Halodule wrightu (shoal grass) dominate and define the South Florida seagrass beds (Zieman 1982, 1990). Thalassic 

is the largest, most abundant, and productive of the South Florida seagrasses. It 

dominates much of the interior of Florida Bay and the bottom of Hawk Channel. 

Syringodium is the numerically dominant seagrass on the western edge of Florida 

Bay and in deeper water and channels with consistent high-velocity conditions . 

Halodule is found in shallow water on bank tops and adjacent to mangrove 

islands. In addition to these three major species, Ruppia maritima, a euryhaline 

angiosperm, is locally abundant in the upper, low-salinity portions of Florida Bay, 

and in the most hypersaline portions of the bay. Three species of Halophila 

(H. decipiens, H. engelmannii, and H. johnsonii) are found primarily around the 

fringes of Florida Bay and on the Southwest Florida continental shelf. Although 

Halophila beds can cover large areas (particularly in deep water), their biomass 

and numerical abundance are low (Continental Shelf Associates, Inc. 1989). 

The same dominant species (except Ruppia) characterize Panamanian 

seagrass beds. The dominant seagrasses in the subtidal Panamanian beds are 

Thalassic testudinum and Syringodium filiforme (Marshall et al. 1993). Halodule is 

found on intertidal areas at the edges of some of the lagoonal seagrass beds 

studied (Marshall, pers. observ.) . Halophila decipiens (and probably other 

Halophila species) are found along the deeper edges of seagrass beds. Thalassic 

also occurs on intertidal reef flats in the study area (Steger and Caldwell 1993). 

Algae 

Associated with South Florida seagrass communities is a diverse group of 

macroalgae and microalgae (Zieman et al . 1989). Seagrass beds possess a 

characteristic assemblage of algal species which can be grouped into three 

categories: 

Those that grow in soft sediments and have either a holdfast or creeping 

rhizoids, such as Halimeda, Penicillus, Rhipocephalus, Udotea, and 

Caulerpa. 

Those that grow on hard substrates, such as Dictyota, Padina, and 

Sargassum. 

15 
Species Composition 

Those that form detached lumps or clumps of free floating drift algae, 

such as Gracilaria, Laurencia, Digenea, and Acanthophora . These all begin 

life on hard substrate, but some fragment and grow as drift algae. 

The microalgal community in South Florida seagrass beds is represented 

almost entirely by epiphytic species growing on seagrass blades . Humm (1964) 

compiled an annotated list of 113 species of algae that were epiphytic on South 

Florida seagrasses. On the west coast of Florida, Rhodophyta (red algae) 

accounted for 45% of the epiphytic species seen. Phaeophytes (brown algae) 

accounted for 12% of the epiphytic species, and Chlorophytes and Cyanophytes 

(green and blue-green algae) each represented 21% of the species seen (Ballantine and Humm 1975). 

Macroalgal genera reported from Panamanian seagrass beds include the 

substrate-attached Halimeda, Penicillus, and Udotea and Dictyota, and the 

filamentous green alga Cladophora. In general, brown algae are sparse in 

Panamanian seagrass beds compared with other tropical areas, and red algae are 

highly seasonal in terms of presence and abundance (Heck 1977). There are 

virtually no data on the epiphytic community of Panamanian seagrasses. 

FAUNA 

There have been numerous faunal studies of South Florida seagrass beds 

(Zieman 1982). Most of the studies have focused on Florida Bay and Biscayne 

Bay; little research has been conducted in shallow seagrass beds surrounding the 

lower Florida Keys (J. Zieman, 1992 pers. comm., Univ. of Virginia). (As noted 

in the previous chapter, the shallow seagrass beds around the lower Florida Keys 

appear most similar in depth range to the Panamanian beds studied). In the 

faunal comparisons below, it is important to recognize faunal variations within 

the South Florida area. The fauna of western Florida Bay is largely southern 

temperate with origins in the Gulf of Mexico, whereas the fauna of the Atlantic 

side of the Keys, or even the bayside lower Keys, is more Caribbean in origin . In 

addition, Florida Bay is a mixing zone that does not have one typical fauna 

(J . Zieman, 1992 pers. comm.) . 

Faunal characteristics of Panamanian seagrass beds are known from studies 

conducted in the spill area before 1986 and from post-spill surveys of unoiled 

beds. All of the pre- and post-spill studies were conducted from the STRI 

Galeta Marine Laboratory, which is located near the Atlantic entrance to the 

Panama Canal and 2.5 km from the spill site (Figure 2). The laboratory opened 

in 1970, and it has been the site of many ecological studies on numerous topics. 

These studies have proven to be an invaluable source of information on pre-spill 

conditions for reef flat habitats . A simple listing of organisms from the 

combined studies of the Galeta Point reef flat habitat includes over 775 species 

(Cubit and Williams 1983). 

16 
Species Composition 

Seagrass bed studies completed before the spill include epifaunal (Heck 1977, 

1979; Weinstein and Heck 1979) and infaunal surveys (Jackson 1973; VasquezMontoya 1979, 1983). In pre-spill studies, epifauna were sampled with an otter 

trawl and infauna were sampled with coring devices. Post-spill studies used 

pushnet collections for epifauna and core samples for infauna (Marshall et al. 

1993) . Various methods of monitoring plant species composition and biomass 

were used in each of these major studies . A similar range of techniques has been 

used to survey plant and animal assemblages in South Florida seagrass beds. 

The following comparisons focus on three main groups for which data are 

sufficient to allow meaningful comparisons: (1) decapods and stomatopods; 

(2) molluscs; and (3) fishes . Species lists for these three groups are presented in 

the Appendix. Table 1 summarizes comparisons at the species and genera levels . 

Decapods and Stomatopods 

Studies of decapod assemblages from seagrass beds are perhaps the best basis 

for faunal comparisons between South Florida and Panama. Heck (1977) used 

an otter trawl, equipped with a 6.3-mm liner, to collect decapod and stomatopod 

crustaceans from seagrass beds within and adjacent to Bahia Las Minas. 

Marshall et al. (1993) collected crustaceans with a pushnet (1.0-mm mesh) from 

several of the same seagrass beds within Bahia Las Minas and in additional 

unoiled seagrass beds that were not included in Heck's study. These two 

collection methods resulted in similar species lists but different relative abundances. Many of the small decapods, including the numerically dominant 

caridean shrimp species were undercollected by Heck's otter trawl, whereas a few 

large, quick-moving decapods may have avoided Marshall's pushnet. The 

combination of methods, however, should have provided a complete description 

of the seagrass bed crustacean fauna. The species listed in Table A.1 are based 

on revisions by Coen and Heck (1983), who reviewed Heck's (1977) collection for 

accuracy. Fewer species are listed in the later paper because specimens were lost 

during the trip from Panama to Tallahassee (K. Heck, pers. comm .) . 

Decapods and stomatopods were collected with throw traps from Florida 

Bay's shallow, mudbank top seagrass beds (Holmquist et al. 1989). Caridean 

shrimp proved to be the numerical dominants in this seagrass bed study. Other 

earlier research on seagrass beds in Florida Bay used a variety of collecting 

techniques and were conducted in areas that included seagrass beds and mud 

bottoms. No similar studies have been carried out in the shallow seagrass beds 

adjacent to the lower Florida Keys or the Atlantic side of the upper Florida 

Keys. 

The decapod and stomatopod faunas of South Florida and Panama share 

numerous species and genera (Table A.1) . Only about 19% of the species occur 

in both areas, but 58% of the species belong to genera that occur in both areas 

(Table 1). Additionally, the two locations share several of the numerically 

dominant species and genera (mostly caridean and penaeid shrimp) (Table 2). 

17 
Table 1 . Summary of fauna) comparisons between seagrass beds of South Florida and Panama. The table summarizes data from 

Appendix Tables A.1 (decapods and stomatopods), A.2 (molluscs), and A.3 (fishes) . 

Species Genera CongenaNc 

Group South Shared South Shared Species 

Florida Panama Total Florida Panama Total 

No. Percent No. Percent No. Percent 

Decapods and 53 66 100 19 19 38 47 61 24 39 58 58 

stomatopods 

00 

Molluscs (total) 166 192 311 47 15 102 125 172 55 32 172 55 

Gastropods 94 136 204 26 13 53 91 109 35 32 114 56 

Bivalves 72 56 107 21 20 49 34 63 20 32 58 54 

Fishes 168 106 224 50 22 116 69 142 43 30 111 50 
Table 2. Numerically abundant caridean and penaeid shrimp from seagrass beds 

of South Florida (Florida Bay--Holmquist et al. 1989) and Panama (Bahia 

Las Minas area--M . Marshall unpubl. data). Numbers indicate rank (n 

each area; for unranked species, 'p' Indicates presence, and '--' Indicates 

absence. 

Rank 

Species 

South Florida Panama 

Thor floridanus 1 -- 

Periclimenes americanus 2 4 

Hippolyte pleuracanthus 3 --* 

Periclimenes longicaudatus 4 10 

Alpheus heterochaelis 5 

Penaeus duorarum 6 5 

Alpheus normanni 7 p+ 

Latreutes fucorum 8 2 

Tozeuma carolinense 9 8 

Leander paulensis 10 -- 

Hippolyte zostericola --* 1 

Thor manningi -- 3 

Alpheidae p+ 6+ 

Latreutes parvulus 13 7 

Sicyonia laevigata -- 9 

* See Gore et al . (1981) for a discussion of the H. pleuracanthus/zostericola complex. 

+ Alpheids from Panama collections have not been identified to species at the time of 

this publication and are reported as Alpheidae, whereas the alpheids reported by 

Holmquist et al. (1989) from Florida Bay were identified to species. Alpheus 

heterochaelis and A. normanni are listed as "present" because they were collected 

during initial alpheid surveys in Panama (M. Marshall, pers. observ.) . 

19 
Species Composition 

Molluscs 

Table A.2 lists mollusc species from seagrass beds of South Florida and 

Panama. The major sources of information on molluscs in Florida Bay (Turney 

and Perkins 1972) and Panama (Racivvin 1969) were studies of living and recently 

dead molluscs in sediments from the two areas. Turney and Perkins (1972) 

described subenvironments within Florida Bay based on molluscan distribution 

patterns . Radwin (1969) studied recent, non-fossil molluscs collected from 

dredge spoil as it was pumped from the shoreward edge of a coral reef in order 

to create the foundation of the refinery (the source of the oil spill) in Bahia Las 

Minas. Tumey and Perkins' species list included living molluscs, whereas 

Radwin's study did not separate living from dead specimens. 

Other molluscan species lists from both South Florida and Panama are from 

general infaunal surveys (Table A.2). The empty shells of dead molluscs, which 

are often the best evidence of the existence of uncommon and rare species in an 

area, were ignored in these other studies. Empty mollusc shells were kept as part 

of the STRI seagrass project, but they have not been sorted (M. Marshall, pets. 

observ.) . 

The species lists suggest that similar molluscan faunas inhabit seagrass beds 

of South Florida and Panama. Although only 15% of the species are common to 

both areas, SS% of the species belong to genera that occur in both areas 

(Table 1). Percentages for gastropods and bivalves are similar. If Radwin's 

(1969) study had been restricted to seagrass bed fauna, the list of species 

reported from Panama would have been shorter, but the degree of overlap at the 

generic and specific levels may have been higher. 

Fishes 

Species lists for fishes in seagrass beds of South Florida and Panama are 

presented in Table A3. Springer and McErlean (1962) sampled seagrass beds at 

Matecumbe Key (Florida Keys) using a seine net. Weinstein and Heck (1979) 

sampled fish by otter trawl in seagrass beds along the Caribbean coast of 

Panama, and also in the Cape Romano area of Southwest Florida. Sogard et al . 

(1989) used throw traps and gill nets to collect fish from seagrass beds on 

mudbank tops within each of the subenvironments of Florida Bay. 

Despite the methodological differences, there are numerous shared and 

oongeneric fish species (Table 1). Although only 22% of the species occur in 

both areas, 50% of the species belong to genera that occur in both areas. 

However, the Panamanian collections include more coral reef-associated species 

than those from South Florida (especially Florida Bay and Southwest Florida) . 

For example, several species of Acanthuridae, Chaetodontidae, Haemulidae, 

Labridae, Lutjanidae, Scaridae, and Serranidae occurred either exclusively in 

Panama or in Panama and the Florida Keys (Springer and McErlean 1962), but 

not in Florida Bay (Sogard et al . 1989) or Southwest Florida (Weinstein and 

20 
Species Composition 

Heck 1979) (Table A.3). Similarities between the Panama and Hawk ChannelFlorida Keys seagrass ichthyofaunas are even more obvious if the species list 

generated for Panama is compared with lists generated by sampling coral reefs 

and adjacent habitats along the Florida reef tract (e.g., Starck 1968; Bohnsack et 

al . 1987). This comparison shows that most (90%) of the species collected in 

Panama also occur in South Florida. Also, only five of the 106 species collected 

in Panama by Weinstein and Heck (1979) are known through taxonomic and 

zoogeographic studies not to occur in South Florida waters . These are Haemulon 

bonariense (Courtenay 1961), Amphichthys hildebrandi (Collette 1966), Rypticus 

brachyrhinus (Courtenay 1967), Serranus flaviventris (Robins and Starck 1961), 

and Diapterus rhombeus (Deckert and Greenfield 1987). Of these, all genera 

except Amphichthys occur in South Florida (Robins and Ray 1986). 

Table 3 compares the most abundant fish species in seagrass beds of 

Southwest Florida (Cape Romano area) and Panama, as determined by 

Weinstein and Heck (1979) using identical methods. The dominant species in 

the two areas are quite different, with reef-associated elements of the tropical 

fauna (e.g., Sparisoma radians) being much less common in Southwest Florida. 

Weinstein and Heck (1979) concluded that the Southwest Florida fish fauna is 

most closely allied with the Carolinean fauna occurring in the Apalachee Bay 

region of Northwest Florida. 

Based on these comparisons, Panamanian fish communities are likely to 

resemble more closely those of seagrass beds in Hawk Channel and the lower 

Keys, rather than Florida Bay. Like the Panamanian beds, the Florida Keys 

seagrass beds exist in oceanic waters and are close to coral reefs (bank reefs or 

patch reefs). Gilmore (1987), in an analysis of zoogeography of western Atlantic 

seagrass fish communities, showed that the waters of Southeast Florida and the 

Florida Keys are inhabited by a tropical fauna, whereas waters of Florida Bay 

and Southwest Florida are inhabited by a warm temperature fauna. This disjunct 

distribution is thought to be primarily due to low winter temperatures on the 

west Florida shelf. 

Other Fauna 

Various other invertebrates were collected during the course of the STRI 

seagrass project (Marshall et al . 1993). Polychaetes, the numerically dominant 

infaunal taxon in Panamanian seagrass beds, have not been completely described 

(K. Fauchald, pers. comm. 1991). Much descriptive work would be required 

before a cross-regional (South Florida to Panama) comparison would be useful at 

the specific level . Despite the earlier works of Vasquez-Montoya (1979) and 

Fauchald (1977), much of this fauna remains poorly known. 

21 
Table 3. Numerically abundant fish species from seagrass beds of South Florida 

(Cape Romano area) and Panama (Bahia Las Minas area) . Collections 

were made with an otter trawl in both areas (Weinstein and Heck 1979) . 

Numbers indicate rank In each area; '--' indicates absence. 

Rank 

Species 

South Florida Panama 

Bairdiella chrysoura 1 - 

Lagodon rhomboides 2 -- 

Eucinostomus gula 3 9 

Haemulon plumieri 4 13 

Orthopristis chrysoptera 5 -- 

Lutjanus synagris 6 3 

Syngnathus scovelli 7 -- 

Chilomycterus schoepfi 8 -- 

Opsanus beta 9 -- 

Eucinostomus argenteus 10 2 

Sparisoma radians -- 1 

Ocyurus chrysurus -- 4 

Sphoeroides spengleri 28 5 

Monacanthus ciliatus 18 6 

Monacanthus setifer 48 7 

Pseudupeneus maculatus -- 8 

Chaetodon capistratus -- 10 

22 
Species Composition 

Echinoderms were extremely rare in Bahia Las Minas seagrass beds from 

September 1986 through August 1989 (end of the seagrass subproject) (Marshall 

et al. 1993). They were seasonally abundant in unoiled seagrass beds outside of 

the immediate vicinity of Bahia Las Minas. Heck (1977) listed six species from 

his otter trawl samples, whereas Vasquez-Montoya (1979) identified eight species 

of echinoderms from within and closely adjacent to Bahia Las Minas. This small 

data base is not sufficient for comparisons with South Florida data. 

Data from other groups also are too limited for meaningful comparisons. A 

small set of isopods, from pushnet and core samples, was identified by 

M. Schotte (National Museum of Natural History, Washington, DC). Amphipods 

were abundant in unoiled areas in pushnet and core samples, but they have not 

been identified. 

23 
CONCLUSIONS 

The purpose of this report was to determine whether and to what extent 

findings from the STRI oil spill study in Panama (Keller and Jackson 1993) can 

be used to predict oil spill impacts upon South Florida seagrass beds. In the 

Introduction, three main factors were cited as affecting the response of seagrass 

beds to oiling : 

Spill circumstances, including spill location, volume, duration, rate of 

release, and chemical composition, effectiveness of spill containment and 

cleanup measures, and unpredictable environmental variables such as 

weather, sea state, and tidal stage at the time of a spill. 

Seagrass distribution and environmental setting, including geographic 

distribution, areal extent, and water depth of seagrass beds; proximity to 

mangrove shorelines; wave energy regime; and history of environmental 

problems. 

Species composition of seagrass bed communities, including both 

seagrasses and associated flora and fauna. 

This report focused on the latter two factors, because spill circumstances 

cannot be predicted in advance. Conclusions regarding the two factors are 

discussed individually below. 

SEAGRASS DISTRIBUTION AND ENVIRONMENTAL SETTING 

Of the environmental factors discussed here, water depth appears to be 

critical in determining effects of oiling on seagrass beds. Intertidal and shallow 

subtidal seagrass beds are the most susceptible to oiling (Zieman et al . 1984; 

Marshall et al . 1990). The most severe effects on seagrasses in Panama occurred 

on intertidal reef flats. Some heavily oiled intertidal beds were killed and may 

never recover (Steger and Caldwell 1993), whereas subtidal seagrass beds 

survived everywhere, with mortality at the margins and more subtle effects within 

the beds (Marshall et al . 1993) . 

In South Florida, seagrasses are ubiquitous, but the largest and most 

productive seagrass beds occur in shallow waters (<1 to 3 m) of Florida Bay and 

in deeper waters (6 to 8 m) of Hawk Channel between the Florida Keys and the 

Florida Reef Tract. The small Panamanian beds, which occur in shallow lagoons 

(<1 m depth) between fringing coral reefs and mangrove shoreline, seem most 

similar to shallow seagrass beds adjacent to islands in the lower Florida Keys. 

They are also similar in depth range to shallow, subtidal seagrass beds on 

mudbanks and around islands in Florida Bay. 

25 
Conclusions 

Because all of the Panamanian seagrass beds studied by Marshall et al. (1993) 

were subtidal, the data could greatly underestimate the severity of oil spill effects 

on Florida Bay seagrasses. Large expanses of seagrass on Florida Bay mudbanks 

are exposed at very low tides and could be highly vulnerable to oiling at certain 

times, similar to the intertidal reef flat beds in Panama studied by Steger and 

Caldwell (1993). The mudbanks are concentrated in the western part of Florida 

Bay, and the shallowest portions of these beds support a seagrass standing crop 

that is twice the bay-wide average (Zieman 1990). 

On the other hand, data from the shallow subtidal Panamanian beds may 

overestimate oil spill impacts on Hawk Channel seagrasses. Because of their 

greater water depth, the Hawk Channel beds might escape oil exposure unless 

passing slicks were sprayed with dispersant to protect "more sensitive" inshore 

habitats. However, under certain conditions (e.g., storms, wave action), oil can 

become mixed into emulsions with seawater and combine with sediment particles 

and sink to the bottom, resulting in oiling of deeper beds (Marshall et al . 1990) . 

Proximity to mangrove shorelines is also an important consideration. All of 

the Panamanian seagrass beds were near mangrove shorelines, resulting in 

chronic exposure over several years as oil leaked out of contaminated sediments. 

Loss of mangroves increased erosion and sedimentation, which also may have 

affected adjacent seagrass beds. Florida Bay is lined by extensive mangrove 

forests which, if oiled, could become a chronic source of oil and sediment 

deposition on Florida Bay seagrass beds. The largest shallow bank beds in 

Florida Bay are well to the south of the mainland mangrove fringe, but 

mangroves also occur throughout the bay on small islands . In contrast, Hawk 

Channel seagrass beds are not surrounded on any side by an oil-absorptive 

mangrove shoreline . 

A key difference between Panama and South Florida is the areal extent of 

seagrass beds. South Florida seagrass beds are extensive, whereas the 

Panamanian beds are small and discontinuous. The difference is important from 

at least two respects. First, despite the size of the Panama spill and the length of 

coastline affected, a relatively small area of seagrass beds was contaminated. 

A much larger area of seagrass beds could be oiled by a single spill reaching 

Florida Bay or the lower Keys. Second, the pattern of recovery from an oil spill 

might be different in South Florida, because some animals could migrate to 

unaffected areas and extensive source beds for new recruits would exist. Small, 

isolated seagrass beds may be subject to severe setbacks or total elimination, as 

contrasted with seagrass beds that are part of a larger, continuous system 

(Zieman et al. 1984) . However, incorporation of oil in seagrass bed sediments 

(as happened in Panama) could inhibit recovery even if new recruits were 

available. 

26 
Conclusions 

SPECIES COMPOSITION 

In general, the species composition of seagrass communities in South Florida 

and Panama is similar. The dominant seagrass species are the same, and 

although many of the epifaunal and infaunal species are different, most species 

belong to genera that occur in both areas. Given the similarities at the generic 

level, there is no basis to assume that these sensitivities differ greatly between the 

two areas. 

However, a distinction must be made between seagrass fauna( communities of 

Florida Bay and those of the Florida Keys. As noted in particular for fishes, 

Florida Bay seagrass fauna have warm temperate amities, whereas those 

associated with the Florida Keys have more tropical Caribbean affinities and are 

therefore more similar to Panamanian seagrass communities . 

Thus, seagrass beds in the lower Keys are most similar to the Panamanian 

beds with respect to water depth, seagrass species, and fauna. In contrast, 

although Florida Bay seagrass beds resemble Panamanian beds with respect to 

water depth and seagrass species, they possess a different (more temperate) 

seagrass fauna. 

SUMMARY 

Data from the STRI study (Kelley and Jackson 1993) could be useful in 

helping to predict effects of an oil spill on South Florida seagrass beds. 

However, environmental and biological differences between the two areas must 

be taken into consideration. 

The seagrass portion of the STRI study (Marshall et al. 1993) focused on 

shallow, subtidal seagrass beds in lagoons between fringing reefs and mangrove 

shorelines. These beds are similar to seagrass beds adjacent to islands in the 

lower Keys with respect to water depth, seagrass species, and associated fauna. 

They are also similar in depth range to extensive, shallow, subtidal seagrass beds 

on mudbanks and around islands in Florida Bay, although Florida Bay possesses 

a more temperate seagrass fauna. The Panamanian beds are in much shallower 

water (<1 m depth) than the extensive seagrass beds (6 to 8 m depth) that occur 

in Hawk Channel between the Florida Keys and the Florida Reef Tract. 

Data from the seagrass study (Marshall et al . 1993) are probably a good 

indication of potential spill effects in shallow, subtidal seagrass beds, such as 

those adjacent to islands in the lower Florida Keys. They are also relevant to 

shallow, subtidal beds in Florida Bay which are similar in water depth and 

proximity to mangrove shorelines. However, the data could greatly 

underestimate the severity of oil spill impacts to seagrasses on Florida Bay 

mudbanks, many of which are exposed during very low tides. A better indicator 

of potential effects on these mudbank seagrasses is provided by the observations 

27 
Conclusions 

of Steger and Caldwell (1993), who documented the apparently irreversible 

destruction of an intertidal seagrass bed at one reef flat site . 

Because of water depth differences, data from the seagrass study (Marshall et 

al . 1993) are less useful for predicting impacts to seagrass beds in deeper waters 

of Hawk Channel. The Hawk Channel beds might escape oil exposure, although 

oil could be deposited under certain circumstances (e .g ., if dispersants were used, 

or if wave action formed oil-in-water emulsions) . Also, unlike the Panamanian 

seagrass beds, the Hawk Channel beds also are not surrounded on any side by an 

oil-absorptive mangrove shoreline. 

The areal extent of seagrass beds is much greater in South Florida than in 

Panama. It is difficult to predict how this difference in scale might affect 

patterns of damage and recovery. Obviously, a much larger area of South 

Florida seagrass beds (and adjacent mangroves) could be oiled by a single spill of 

similar size. Patterns of recovery might be different in South Florida because 

some animals could migrate to unaffected areas and extensive source beds for 

new recruits would exist. However, chronic oil release from contaminated 

mangrove sediments and incorporation of oil into seagrass bed sediments (as 

happened in Panama) could inhibit recovery even if new recruits were available . 

28 
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33 
APPENDIX 

SPECIES LISTS 

A-1 
Table A.1 . Comparison of decapod and stomatopod crustacean species lute for 

South Florida (Florida Bay) and Panama (Bahia Las Minas area). 

Source codes: for Florida, (1) Holmquist et al. (1989) ; for Panama, 

(2) Heck (1977) as modified in Coon and Heck (1983), and (3) Marshall 

(unpublished data). 

South 

Species Florida Panama 

DECAPODA 

Panaaidaa 

Metapenaeopsis msrtinella 2,3 

Penaeus duorarum noGialis 1 2,3 

Trackyp enaeus similis 3 

Sicyonidao 

Sicyonis laevigata 2,3 

Stanopodidae 

Stenopus hispidus 2 

Palaemonidae 

Lesnder tenuicornis 1 2,3 

L. paulensis 1 

Palaemon floridanus 1 2 

P. northropi 3 

Periclimenes americanus 1 2,3 

P. iridescens 1 

P. longicaudatus 1 3 

Gnathophyllidae 

Gnathophylloides mined 2,3 

GnethopAyllum americanum 3 

Procassidae 

Ambidexter symmeficus 1 2,3 

Process fimbrieta 2,3 

P. bermudensis 1 

A1pheidae 

Alpheus armillatus 1 2 

A floridanus 2,3 

A formosus 2 

A normanni 1 2 

A heterochaelis t 

Synalpheus fritzmulleri 2,3 

S. goodei 2 

S. pandionensis 2 

S. townsendi 2,3 

Ogyrididas 

Ogyrides alphaerostris 1 

HiPpolytidae 

Hippo lyte zostericola 2,3 

H. pleuracanthus 1 

Labeutes parvulus 1 3 

L. fucorum 1 2,3 

Thor manningi 2,3 

T. dobkini 1 

T. floridanus t 

Tozeuma carolinense 1 2,3 

Tiachycaris restrictus 2,3 

Scyllaridae 

ScyIlarus sp. 2 

Palinuridae 

Palinurus argus 1 2,3 

Diogenidae 

Celcinus tibicen 2 

Clibanarius antillensis 2,3 

Dardanus venosus 2 

Paguristes limonensis 2 

P. anomalus i 

P. tortegae 1 

Paguridaa 

Pa gurus bonairensis 2 

P. brevidactylus 2 

P. mclaughlinae 1 

A-3 
Table A.1 . (continued). 

Species 

South 

Florida Panama 

Porcollanidaa 

Megalobrachium mortenseni 

Pebolishtes armatus 1 

P. galethinus 

Dromiidao 

Dromidie antillensis 

Calappidaa 

Ca/apps angusta 

C. ocellata 

Cala S P 

He ~ udibundus 

Ma Idao 

Chorinus hems 

Libinis dubis 1 

Metaporhapsis calcsrata 1 

Macrocoeloms diplacanthum 

Microphrys bicomutus 1 

Mithrax forceps 1 

M. ruber 

Pelia mubca 1 

Pitho aculeata 

P. Iherminieri 

P. quadridentata 

P. sexdentata 

P. anisodon 1 

Podochela gracilipes 

P. nisei 1 

P. sidneyi 

Sterarhynchus seticomis 

Parthenopidae 

Heterocryp to granulate 

Goneplacidae 

Cyrtoplax spinidentafa 

Eucrafopsis crassimenus t 

Portunidaa 

Callinectes omatus 1 

C. sepidus t 

C. dance 

Cronius Tuber 

C. dmidulus 

Portunus depressifrons t 

P. gibbesii 1 

P. ordwayi 1 

P. sayi 1 

P. sebae 1 

P. spinimanus 1 

Xanthidae 

Menippe mercensria 1 

Dyspanopeus texanus 1 

Eurypanopeus depressus 1 

Micropanope sp. 1 

Neopanope packardii 1 

Panopeus occidentalis 1 

P. simpsoni 1 

Pilumnus desypodus 

Rithropanopeus harrisii 1 

Pinnotheridae 

Pinnixia sayana 1 

Ocypodidaa 

Uce SP. 1 

STOMATOPODA 

Squilltdae 

Meiosquilla quadridens 1 

Gonodactylidae 

Gonodnctylus lacunstus 

Psaudosqulllidaa 

Pseudosquilla ciliate 1 

2 

2 

2 

2 

2 

2 

2,3 

2 

2,3 

2 

2 

2 

2,3 

2 

2 

2 

2 

2,3 

2 

2 

2 

2 

2,3 

2,3 

2 

2 

2 

2 

2 

2 

A-4 
Table A.2. Comparison of mollusc species lists for South Florida (mainly Florida 

Bay) and Panama (Bahia Las Minas area). Numerous other 

Panamanian species, not included in this list because of unknown 

habitat associations, were recorded (Radwin 1969) from the dredged 

material used as a foundation for the Bahia Las Minas oil refinery 

(the source of the 1986 spill) . Source codes: for Florida, (1) Tabb 

and Manning (1961), (2) Tabb et al. (1962), (3) Hudson et al. (1970), 

(4) Turney and Perkins (1972), and (5) McClanahan (unpublished 

data) ; for Panama, (6) Radwin (1969), (7) Jackson (1973), (8) Heck 

(1977), (9) Vasquez-Montoya (1979), (10) Cubit and Williams (1983), 

and (11) Marshall and Batiste (unpublished data). 

South 

Species Florida Panama 

GASTROPODA 

Fissurellidao 

Emarginula pumila 6 

Hemitoma octoradiata 6 

Diodora caynensis 1,3 6,11 

D. mete 1 

D. listen 

D. dysoni 2 

Diodors 

SP. 

Lucapina suffuse 6 

Fissurella lescicularis 6 

F. nodosa 

Acmaaidae 

Acmeee pusfulata t 1 

Trochidae 

Calliostoma jujubinum tampaense 1,2,3,4 

C. euglyptum 6 

C. javanicum 6 

Cittarium pica 6 

Tegula /asciata 1,2,3,4,5 6 

Soleriorbis schumoi 6 

S. corylus 6 

S. hondurasensis 6 

S. infracarinsta 6 

S. shimeri 6 

Armclimax schumoi 6 

Turbinidae 

Turbo castaneus 1,2,3,4 6 

T. canaliculetus 5 

T. lilosus 6 

Astreea phoebie 3,5,5 6 

A tecta americana 3,4,5 6 

A caelata 6 

A longispina 4 

Liotia ficarinata 6 

Neritidaa 

Smsragedia viridis 4 6,11 

Neritina virginea 6,11 

Architectonicidae 

Architectonics nobilis 6 

Heliacus parried 6 

Littorinidaa 

LiKorina angulifera t 

Littorina sp. 6 

Vermetidaa 

Vermicularia sp. 1 

Vermetus varians 6 

Aletes mcgintyi 6 

A nebulosus 6 

A lloridanus 6 

Stephopoma myrakeenee 6 

Spiroglyphus ennuletus 6 

Thylacoides sp. 6 

A-5 
Table A.2. (continued) . 

Species 

South 

Florida Panama 

Modulidaa 

Modules modules 1,2,3,4,5 6,8 

M. carchedonius 6 

Potamididaa 

Cerithidea costats 1 

C. scalariformis 1 

Carkhidaa 

Bstillana minima 1,3 

Cerithium floridanum 1,2,5 

C. muscarum 1,3,4,5 

C. ebumeum 1,2,4,5 8,9,11 

C. algicola 2 6 

C. litteratum 4,5 6,11 

C. moenense 6 

C. variabile 4 6,9,11 

Bittium varium 1,4 6 

Alabina cerithioides 6 

Litiopa melanostoma 6 

Epitoniidae 

Epitonium sp. 1 

E. folieceicostum 6 

Opalia crenats 6 

Depressiscala nautilae 6 

Calyptraeidaa 

Calyptraea centrelis 1,2 6 

Crucibulum auriculum 6 

Crepidula maculosa 6 

C. cornexe 1,3 6 

C. aculeata 1 

C. plane 1,2,3 6 

Crepidule sp. 4 

Strombidae 

Strombus elates 1 

S. gigas 5 

S. pugilis 6 

S. rsnius 5 6,11 

Eratoidaa 

Erato maugeriae 1,2 6 

Trivia candidula 1,2 

T. pediculus 6 

T. quadripunctata 6 

Trivia sp. 4 

Cypraeidae 

Cypraea cervus 5 

Naticidae 

Polinices duplicates 1,2 

P. lacteus 6,11 

P. hepaticus 11 

Neticea canrena 1 6 

N. livide 2 

Tectonaticea pusilla 6 

Sinum maculatum 6 

G/ypheithema floridane 6 

Tuibinellidae 

Vesum muricatum 11 

Cassiidaa 

Phalium granulatum 1 

Ficidae 

Ficus communis 1 

Muricidaa 

Murex recurvirostris rubidus 1,2 8 

M. ponum 1,2 

M. florifei 1,2 

M. cellulosus 1,2,3 

M. brevifrons 4 

M. rubidus 6 

M. woodringi 6 

Chicoreus florifer 6 

A-6 
Table A.2. (continued). 

South 

Species Florida Panama 

Muricidaa (continued) 

Muricopsis ostrearum 1,3 

M. oxytetus 6 

M. philippina 6 

Risomurex schrammi 6 

R. muricoides 6 

MuNcidaa (continued) 

Monrla (Drupe) noduiosa 5 6 

M. didyma 6 

Theis deliuides 6 

Urosalpinx tampaensis 1 

Eupleura sWcidentata 1,3 

E. caudate 2 

Columbellidae 

Columbella mercaroria 1,2,4,5 6 

C. rus8coides 1,2,3,4 

Anachis avara 1,2,4 

A obese 1,2 6 

A translirata 1,2 

A pebi 6 

Astyris lunata 6 

Conella ovuloides 6 

Cosmioconcha nifens 6 

Nitidella nitida 6 

Steironepion monilifera 6 

Za/rona pulchella 6 

Mibella luneta 4 

Buccinidaa 

Engina turbinella 6 

Barlya intricate 6 

Ceducifer edelus 6 

Cantharus tinchrs 1,5 

C. auritulus 6 

Pisanis pusio 6 

Melongenidae 

Melongena corona 1,3,4 

M. melongena 6 

Busycon contrarium 1,2,3 

B. spirafum 1,2,3 

Nassariidae 

Nassarius vibex 1,3,4 9 

N. ambiguus 2,4 

N. cinisculus 6 

Fasciolariidae 

Fasciolaria tulips 1,2,3,4,5 6,8,9,11 

F. hunteria 1,2,3 

Fusilahius cayohuesonicus 6 

Pleuroploca gigantea 1,2,4,5 

LaGirus infundibulum 6 

L. carinifera 6 

Leucozonia mass 6 

Olividae 

Olive caribaeenis 6 

Olivella nivea 6 

O. chiriquiensis 6 

O. sp. 1 

O. minute 3 

Minioliva myrmecoon 6 

Margfnellidae 4 

Marginella denticulate 5 

M. ousts 

M. fPrunum) spicine 

5 

1,3,5 

P. cameum 6 

Volvarina evens 6 

Gibberuls bocasensis 6 

G. ovuliformis 6 

A-7 
Table A.2. (continued). 

Species 

South 

Florida Panama 

Conidaa 

Cones spurius spurius 6 

C. regius 6 

C.deucus 6 

C. grenulatus 6 

C. lugillierti 6 

C. pygmaes 6 

C. mus 6 

C. mindanus 6 

C. floridenus 1 

C. stearnsi 1,2,3 

C. jsspideus 1,2 6 

Tarebridae 

Terebra dislocate 1,2 

T. taurinum 6 

T. spei 6 

Turridae 

Polyslira albida 6 

Drillis albinodeta 6 

D. albomaculata 6 

Syntomodrillie lissotropis 6 

Crassispira auberti 6 

C. harfordiana 6 

C. chazaliei 6 

C. ebeninn 1,2 

C. osbearum 1,2 

C. albomaculata~ 2 

Cerodrillis then 1 ,3 

C. perryae 2 

Nannodiella oxylata 6 

Brachycythars biconica 6 

Ithycylhara psila 6 

Mengelia Now 6 

M. stellata§ 4 

Mangelie sp. 4 

Thelecythara floridana 6 

Acmaturris sp. 6 

Daphnella lymnaeiformis 6 

Bullidae 

Bulls occidentalis 6 

B. striate 1,3 

Atydae 

Atys guildingi 6 

a riisiena s 

A caribaea 1 

Haminoea elegans 1 

H. succinea 1 

H. antillarum 1,3 9 

Aplysidaa 

Bursatelle leschi plei 1 

Plourobranchidae 

Pleurobranchus aHanticus 1,2 

Ellobiidae 

Melampus coffees 1 

Ischnochitonidaa 

Chaetopleura apiculata 1,2 

Ischnochiton papillosus 3 

PELECYPODA 

Arcs zebra 6 

A imbricata 6 

Arcopsis adamsi 1,2,3,4 6,9,11 

Anadara chemnitzi 6 

A notebilis 1 6,11 

Barbetie cancellaria 6 

B. candide 6 

NoeBa ponderosa t 

A-8 
Table A.2 . (continued). 

Species 

South 

Florida Panama 

G cymoridao 

G ~ jncym eris pectinata 

M i id 

1,2,3,4 6 

yt l ao 

Brechiodontes exushrs 1,3,4 7 

8. recurvus 1 

Amygdalum papyna 1 

Musculus lateralis 1,2,3 

Modiolus americanus* 4 6 

Litho hags antillerum P 6 

Botu fusca 6 

Ptariidaa 

Plane colymbus 1 

Pinnidae 

Pinna camea 1 

Atrina rigida 1,2,3,4 

A serrate 1,2 

Pactinidaa 

Pecten ziczac 1,2 

Aequipecten muscosus 1,2 

A irradians 1,3 

Pecfen gibbus 1 

Llmidae 

Lima pellucida 1,3,4 

Anomiidaa 

Anomie simplex 1 

PteNidae 

Pincteda radials 3,4 

Ostreidaa 

Ostres equestris 1 

Crassostrea virginica 1 

Carditidae 

Cardita floridane 1,2,3,4 

Venericerdia tridentata 1 9 

Lucinidae 

Lucina pensylvenica 6 

L. nessula 6 

L. pecfinatus 6 

L. muricatus 6 

L. amiantus 1,2 

L. multilineste 4 

Phacoides pectinatus 1 7 

P. nassula 1,4 

Codekia orbiculata 1,2,3,4 6,9 

C. orbicularis 4 6,9,11 

C. pectinella 6 

Ungulinidae 

Diplodonta punctata 11 

Cardiidae 

Trachycardium isocardie 6 

T. muncatum 1,2,4 6 

T. egmon6anum 1,2 

Laevicardium laevigatum 4 6 

L. morioni 1,3,4 6 

L. multilineata 6 

Trigonocarclia antillarum 6 

Americerdia media 6 

Papyridea soleniformis 6 

Vanaridaa 

Chione cancellats 1,2,3,4 6,9,11 

C. paphia 6 

Anomalocardia cunimeris 1,3,4 

Pitsr simpsoni 1 

P. fulminate 4 

Transenella cubaiana 3 

T. s0impsoni 3 

Tiansenelle sp. 4 

Macrocellista nimbosa 1 

M. maculate 1 6 

A-9 
Table A.2. (continued). 

Species 

South 

Florida Panama 

VanoNdaa (continued) 

Dosinia elegens 1 

D. discus 

Calbcaidia albida 6 

Aidgona listen 

Gouldia cerise 6 

Tolltnidaa 

Telling listen 

T. angulosa 6 

T. martinicensis 

T. versicolor 1,2 9 

T. promera 1 

T. tamQaensis 1,3 

T. similis 1,3,4 

T. lineata 1,3 

T. alternate 1 7,11 

T. punicea 1 6 

T. texana 4 

T. mars 4 

Tellidora cristata 1,2 

Scissula exilis 6 

Macoma mitchili 1 

M. consbicta 1 9 

M. lento 

Arcopagis fausta 6 

Sartwlidae 

Semele proficua 1,2 6 

S. purpurascens 6 

Abra aequalis 1 

A lioica 

Cumingia tellinoides 4 

Sanguinolarildae 

Tagelus plebius 1 

T. divisus 1 11 

Mactridaa 

Spisula solidissima 1 

Lsbiosa plicatella 1 

Mac" fragilis 1,2,4 

Corbiculiidae 

Polymesoda caroliniana 1 

Pseudocyrena mariHma 4 

Corbulidae 

Corbula contracts 1,2 

C. banattiana 1,2 

C. swiftiana 2 6 

C. dietziana 6 

C. cubaniana 6 

C. caribaea 6 

Notocorbula operculata 6 

Pholadidaa 

Cyriopleura costata 1 

Lyonsiidae 

Lyvnsia hyaline 

Cuspidariidae 

1,3,4 

Cardiomya costellats 4 

listed in (2) as Monilis ira albinodonta . 

+ listed in (4) as VolselP americana . 

§ listed in (4) as Stellatoma stellata . 

A-10 
Table A.3. Comparison of fish species lists for South Florida and Panama. 

Collection methods used to each study are discussed in the text. 

Source codes: for South Florida, (1) Springer and McErlean 

(1962)--Matecumbe Key; (2) Weinstein and Heck (1979)--Cape 

Romano area; and (3) Sogard et al. (1989)--Florida Bay; for Panama, 

(2) Weinstein and Heck (1979)--Bahia Las Minas area. 

South 

Species Florida Panama 

Rhincodontidaa 

Gfnglymostoma cirretum 

Carcharhinidae 

Carcharhinus limbatus 3 

Negaprion brevirostris 3 

Sphyrnidao 

Sphyma tiburo 3 

Pristidaa 

Pristis pectinata 3 

Rhinobatidae 

Rhinobatos len8ginosus 3 

Torpedinidae 

Narcine brasiliensis 2 

Dasyatidaa 

Dasyabs sabina 2 

Urolophidae 

Urolophus jamaicensis 

Elopidao 

Elops saunrs 1,3 

Atbulidaa 

Albula vulpes 3 

Muraanidae 

Gym nothorax vicious 

Ophichthtdae 

Ahlia egmornis 3 

INyrophis punctatus 3 

Clupoidaa 

Hsrengula jeguana 1,3 

Jenkinsia lamprotsenia 1,3 

Opisthonema oglinum 1,3 

Sardinella aunts 1 

S. brasiliensis 3 

Sardinella sp. 

Engraulidae 

Anchoa cubana 1 

A mitchilli 1,3 

A lyolepis 

A hepsetus 3 

Anchovielle perfasciste 1 

Synodontidae 

Syradus foetens 1,2,3 

S. poeyi 

S. intermedius 

Arildaa 

Anus fells 2,3 

Bagre merinus 1,3 

Batrachoididae 

Opsanus beta 1,2,3 

Amphichthys hildebrandi 

Porichthys plecfrodon 3 

Gobieaocidae 

Acyrtops beryllinus 1 

Gobiesox strumosus 1 

Antennariidae 

Hisfrio histrio 1,3 

Antennarius scaber 

Ogcxephaiidae 

Ogcocephalus radiates 2 

Bythitidae 

Ogilbia cayorum 3 

A-11 
Table A.3. (continued). 

South 

Species Florida 

Carapidae 

Cuapus bermudensis 1 

Exocoatidao 

Chiodorus atherinoides 3 

Hemiramphus brasiliensis 1,3 

F1yporhamphus unifasciatus 3 

Bafonidaa 

Strongylura marina 3 

S. notata 1,3 

S. timucu 1,3 

Tylosurus crocodilus 3 

Cyprinodontidae 

Adinia xenica 3 

Cyprinodon variegates 

Flo idi hth i 

3 

r c ys csrp o 1 3 

Fundulus sp. 

, 

3 

Lucanis parva 1,3 

Poecilidae 

Poecilia ladpinne 3 

Athorinidao 

Atherinomorus stipes 1,3 

Hypo!ttherins harringtonensis 1,3 

Menidia peninsulee 3 

Menidia sp. 

Aulostomidaa 

Aulostomus maculates 

Fistulariidao 

FistWsria tsbacsrie 

Holocentridae 

Holocentrus rufus 

Syngnathidaa 

Anarchopterus criniger 3 

Cosmocampus albirostris 1,3 

C. brachycephalus 1 

Hippocampus erectus 2 

H. reidi 2 

H. zosterae 1,2,3 

Syngnathus dunckeri 1 

S. floridae 1,2,3 

S. louisianae 1,2,3 

S. scovelli 1,2,3 

Micrognathus crinigerus 2 

Cantropomidae 

Centropomus pectinatus 3 

Sarranidao 

Diplectrum bivittatum 3 

D. formosum 2 

Epinephelus sbiatus 

E. mono 2 

E. itajara 

Alphestes afar 

Aypticus brachyrhinus 

R. saponaceus 

Myctoperca bonaci 1 

M. microlepis 2 

Serrenus flaviventris 

S. subligarius 2 

Serraniculus pumilio 2 

Apogonidao 

Asbapogon elutes 

Phaeoptyx pigmentaria 

Pomatomidae 

Pomatomus saltatrix 3 

Rachycentridae 

Rachycentron canadum 3 

Echeneidae 

Echeneis naucrates t 

Panama 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

A-12 
Table A.3. (continued) . 

South 

Species Florida Panama 

Carangidaa 

Trachinotus cerolinus 3 

T. falcatus 3 

Car+anx crysos 3 

C. hippos 1,3 

C. baKholornaei 2 

C. latus 3 2 

Chloroscombrus chrysurus 2 

Oligoplites saurus 3 

Selene wmer 3 

Lutlanidaa 

Lutjanus synegris 2,3 2 

L. griseus 1,2,3 2 

L. enslis 1 2 

L apodus 1 2 

L. jocu 2 

Ocyurus chrysurus 1 2 

Lobotidao 

Lobotes surinamensis 1 

Gorreidaa 

Eucinostumus ergenteus 1,2 2 

E.g ula 1,2,3 2 

E. lelroyi 1 2 

E. harengulus 3 

Eucinostomus spp. 3 

Diapterus rhombeus 2 

Games cinereus 3 2 

Haemulidaa 

Haemulon parre 1,3 2 

H. plumieri 1,2,3 2 

H. sciurus 1,3 2 

H. nurolineatum 1,2,3 2 

H. boneriense 2 

H. flawlineatum 1 2 

H. chrysergyreum 3 

Haemulon sp. 3 

Anisotremus virginicus 2 

Orthopristis chrysoptera 1,2,3 

Sparidae 

Archosargus probatocephalus 3 

A rhomboidalis 3 2 

Lagodon rhomboides 1,2,3 

Celamus arctifrans 1 

C. penny 2 

Sclaanidaa 

Baiidiella batebana 2 

B. chrysours 2,3 

B. sanctaeluciae 2 

Cjmoscion nebulosus 2,9 

C. arenarius 3 

Leiostomus xanthurus 3 

Menticirrhus saxatlis 1 

Poqonias cromis 3 

Sciaeraps ocellatus 3 

Odontoscion dentex 2 

Mullidaa 

Pseudupeneus maculatus 1 2 

Ephippidao 

Chsetodipterus faber 1,2,3 2 

Chaetodontidae 

Cheetodon capistratus 2 

C. strietus 2 

C. ocellatus 2 

Pomacanthidae 

Pomscanthus pare 2 

A-13 
Table A.3. (continued). 

Species 

South 

Florida Panama 

Pomacantridaa 

PomacenErus leucostictus t 

P. plerrifrons 

Labridaa 

Dorafonatus megalepis 1 

Halichoeres biviKetus 1 

H. poeyi 

Hemipteronotus martinicensis t 

Scaridaa 

Spaiisoma radians 1 

S. chrysopterum 1 

S. iubripinne 1 

S. wide 1 

Sparisoma sp. 3 

Cryptotomus roseus 

Nicholsins usta 1,2 

Scarus croicensis 1 

S. guscamaia 1 

Mugilidae 

Mugil cephalus 3 

M. curems 1,3 

M. gyrens 3 

M. trichodon 1 

Sphyraanidae 

Sphyraena barracuda 1,3 

S. guachancho 

S. picudilla 

Polynemidaa 

Polydactylus virginicus 

Cltnidaa 

Chaenopsis ocellate 3 

Paraclinus fasciatus 3 

P. marmoratus 1,3 

Labrisomus nuchipinnus 

Malacoctenus aurolineatus 

M. macropus 1 

Blannidaa 

Chssmodes saburrae 3 

Hypsoblennius ionUhas 2 

Cailionymidae 

Diplogrsmmus pauciradiatus 1,3 

Gobi(dae 

Barbulifer ceuthoecus 1,3 

Gobionellus hastatus 

G. saepepallens 3 

G. stigmaturus 1 

Gobiosome robustum 1,2,3 

Microgobius gulosus 

C h l f 

3 

oryp opterus g auco raenum 

Bathygobius curacao 

Acanthuridaa 

Acanthurus chirurgus 1 

A bahienus 1 

Dactyloptaridae 

Dactylopteros volitans 

Stromatatdaa 

Nomeus gronovii 3 

Scorpaanidae 

Scorpaena bergi 

S. brasiliensis 1 

S. grandicomis 1 

S. isthmensis 

S. plumieri 1 

Triglidae 

Prionotus salmonicolor 1 

P. scitulus 2 

P. tribufus 1 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

A-14 
Table A.3. (continued) . 

South 

Species Florida Panama 

Bothidaa 

Mcyclopsetta quadrocellata 2 

Citharichfhys macrops 1 

C. spilopterus 2 

Bothus ocellatus 1 2 

B. meculiterus 2 

Syacium micrurum 2 

Paralich1hys albigutta 2,3 

Ebopus crossotus 2 

Solaidaa 

Achirus lineatus 1,3 

Trinectes inscriptus 1 

T. meculstus 3 

Cynoglossidae 

Symphurus plagiusa 1,3 2 

Balistidaa 

Balistes capriscus 1 

Monacanihus ciliafus 1,2,3 2 

M. hispidus 1,2,3 

M. sedfer 2 2 

M. tucked 2 

Aleuterus schoepfi 2 

Ostraclidae 

Lactophrys quedricomis 1,2,3 2 

L. triqueter 2 

L. trigonus 1,2 2 

Totraodontidae 

Sphoeroides nephelus 1,2,3 2 

S. testudineus 2 

S. spengleri 1,2,3 2 

Carrthigester rostrata 2 

Dlodontidaa 

Chilomycferus schoepfi 1,2,3 

C. aMennatus 2 

Diodon holocenthus 1 2 

A-15 
Q1l4 NT F y~ 

'st F 

~ y The Department of the Interior Mission 

o`' ~ p As the Nation's principal conservation agency, the Department of the Interior has responsibility for 

most of our nationally owned public lands and natural resources. This includes fostering sound use 

of our land and water resources; protecting our fish, wildlife, and biological diversity ; preserving the 

_ ~ 

~, 

environmental and cultural values of our national parks and historical places; and providing for the 

ACH 9 enjoyment of life through outdoor recreation . The Department assesses our energy and mineral 

resources and works to ensure that their development is in the best interests of all our people by 

encouraging stewardship and citizen participation in their care. The Department also has a major 

responsibility for American Indian reservation communities and for people who live in island territories 

under U.S. administration . 

The Minerals Management Service Mission 

As a bureau of the Department of the Interior , the Minerals Management Service's (MMS) primary 

responsibilities are to manage the mineral resources located on the Nation's Outer Continental Shelf 

(OCS), collect revenue from the Federal OCS and onshore Federal and Indian lands, and distribute 

s 

those revenues. c4 

Moreover, in working to meet its responsibilities, the Offshore Minerals Management Program 

administers the OCS competitive leasing program and oversees the safe and environmentally sound 

exploration and production of our Nation's offshore natural gas, oil and other mineral resources. The 

MMS Royalty Management Program meets its responsibilities by ensuring the efficient, timely and 

accurate collection and disbursement of revenue from mineral leasing and production due to Indian 

tribes and allottees, States and the U.S . Treasury . 

The MMS strives to fulfill its responsibilities through the general guiding principles of: (1) being 

responsive to the public's concerns and interests by maintaining a dialogue with all potentially affected 

parties and (2) carrying out its programs with an emphasis on working to enhance the quality of life for 

all Americans by lending MMS assistance and expertise to economic development and environmental 

protection.