Literature values for mean number per platform (MN), size range at platform (SR, cm FL), mean length of individuals observed (ML, cm FL), age range (AR yrs), and biomass per platform (B kg wet wt) for five abundant species of fishes collected from Gulf of Mexico oil and gas platforms.
Abstract
There are over 2300 standing oil and gas platforms in the northern Gulf of Mexico (GOM). It has been argued that platforms provide reef-like habitat that increases the growth and survival rates of fishes by increasing prey availability and affording shelter for protection from predators, provide additional spawning substrate, and by acting as a visual attractant for organisms not otherwise dependent upon hard bottom. Platforms differ from most natural habitats, and from traditional artificial reefs, in that their vertical profile extends upward through the water column into the photic zone and the sea surface. Increased habitat quality on, or immediately around, oil and gas platforms are thought to be derived from increased in situ food production associated with encrustation by fouling organisms. In this chapter, we address the issue of how to evaluate the role of artificial reefs by first establishing levels of evaluation for individual fish species found on oil and gas platforms in the GOM. The levels of evaluation relate to the amount and adequacy of the available information, which was populated with an extensive literature and data search. Three levels of assessment are established, analogous to the levels of analysis established National Oceanographic and Atmospheric Administration (NOAA) Fisheries for identification of Essential Fish Habitat. More than 1300 documents, including reports, stock assessments, other gray literature, and papers published in the primary literature, were used to complete this chapter. When available, published literature was the preferred source of information.
Keywords
- Gulf of Mexico
- oil and gas platforms
- fish
- fisheries
- biomass production
1. Introduction
In the waters of the northern Gulf of Mexico (GOM) there are over 2300 standing oil and gas platforms that together constitute the largest
Artificial reefs, such as oil and gas platforms, may be useful tools for fishery managers if they increase reef fish biomass production, but many researchers question whether or not they are a positive influence on fish stock dynamics. If artificial reefs constitute habitat that is otherwise limiting for reef-associated fishes, then they may be viable management tools. If they are simply attracting fish, then they may be promoting overfishing. Unfortunately, the extent to which structures have influenced the status of exploited fish stocks, either directly via population production rates or indirectly through changes in fishing mortality rates, is still not well understood. The structures may alter fish populations and communities as a result of altering ecosystem structure and function. The effects on exploited fish stocks could also be detrimental if the high levels of fishing mortality rates do not result in compensatory processes that lead to increases in stock production.
2. Methods
In this chapter, we evaluate the role of artificial reefs by first establishing levels of evaluation for individual fish species found on oil and gas platforms in the GOM. The levels of evaluation are dependent upon the amount and adequacy of the available information, which was populated with an extensive literature and data search. Three levels of assessment are established, analogous to the levels of analysis established National Marine Fisheries Service for identification of Essential Fish Habitat (EFH). More than 1300 documents, including reports, stock assessments, other gray literature, and papers published in the primary literature, were used to complete this chapter. When available, published literature was the preferred source of information.
Level 1–For species about which little process information is known, the evaluation is based simply upon whether the species has been observed in association with platforms or artificial reefs.
Level 2–Here we used the conceptual model of Bohnsack [1] (qualitative, Figure 2). This conceptual model centers on the attraction vs. production issue, which encompasses much of the debate about the ecological role of artificial reefs (including oil and gas platforms) in a complex and dynamic coastal geography. The difference between Level 1 and Level 2 assessments is the degree of inference at the process level about the species in question, even if the process information (e.g., fishing mortality, site fidelity) is poorly documented. As such, relative knowledge of where a species falls along the continuum of data availability and confidence for several process-related variables provides significant insight into how that species may be affected by platforms. The evaluation presented here is based upon information reported In FishBase® (http://www.fishbase.org/), combined with expert opinion, and is used to provide relative species-specific assessments of: (1) Site fidelity (High, Moderate, Low); (2) Whether or not a directed fishery exists for this species (Yes or No, includes recreational fishing); (3) Whether diet is derived directly from reef habitat (Reef, Benthic, Pelagic); (4) Whether population size is believed to be limited by recruitment or habitat limitation (Habitat limited, Recruitment limited); (5) Type of behavior of adults (Reef, Demersal, Pelagic, Highly Migratory); and, (6) A summary judgment about whether the species is reef or habitat dependent, and the type of habitat on which some dependent species are most often found (e.g., Sargassum, Sea Grass, Hard Bottom).
Level 3–Based upon our extensive review of the literature, data for only five species of fishes were deemed sufficient for more complex analysis of estimating production; these are red snapper
The first model used in Level 3 evaluations is the semi-quantitative model described in Powers and colleagues [2], which uses the species-specific fish biomass production of a population on a reef (here a platform) weighted by the degree to which growth (biomass production) is attributable to prey resources produced on the reef. The production estimate for each species is multiplied by an index of reef exclusivity (IRE) derived from quantitative diet data. Applying the IRE, annual production (P) of a species attributed to a platform (AP; kg platform-1 yr-1) is calculated by:
where AP is a measure of relative species-specific production attributable to a platform. In the original equation, there was a term for the difference between pre-structure and post-structure biomass, but we are unable to provide pre-structure data because almost all of the platforms on the continental shelf were employed more than 25 years ago. We also believe that an estimate of biomass on a seafloor lacking structure, when compared to biomass after a structure has been constructed, may result in an overestimate of new biomass production because of the likelihood that individuals of many species are simply attracted to standing platforms.
The second method of estimating annual production for Level 3 is an empirical model [3]:
where P is production (g dw d-1), B is biomass (kg), and T is temperature (°C). We assumed that g dw = g wet weight * 0.20. The data used to estimate B (g) is based upon data obtained from published literature on biomass, estimated daily somatic production, and ambient water temperature for 62 species of fishes collected from numerous locations in Australia and elsewhere. Temperature data were summarized from several Gulf of Mexico studies on the continental shelf.
The previous and following methods of estimating production require an estimate of biomass on a platform. To make this estimate, we first calculated the simple arithmetic mean number of fish by species on a platform by summing all of the available estimates of numbers observed, based mostly upon visual surveys using scuba diving. In addition to numbers of individuals, length ranges (cm) are also reported for each species.
The third estimate of production for Level 3 is based upon methods described in by Ricker [4] where annual production is estimated by:
where
Where
Species | MN | SR | ML | AR | B (kg) |
---|---|---|---|---|---|
Red snapper | 1884 (range 905–4632) | 25.5–79.1 | 295.3 | 2–10 | 886 |
Bluefish | 1438 (range 282–4000) | 45–50 | 475 | 1–6 | 1489 |
Atlantic spadefish | 4177 (range 10 –5323) | 10–50 | 30.0 | 1–8 | 2618 |
Sheepshead | 2250 (range 150–17,000) | 22–50 | 360 | 2–5 | 1774 |
Blue runner | 6260 (range 427–25,188) | 30–36 | 33.5 | 2–6 | 4152 |
Species | Tmax | G | M | AD | L• | K | t0 | a | b | Source |
---|---|---|---|---|---|---|---|---|---|---|
Red snapper | 57 | 0.31 (at Tmax G=0.05) |
0.10 (at Tmax M=0.07) |
2–10 | 94.1 | 0.18 | –0.55 | 0.0165 | 3.03 | [5–7] |
Blue runner | 11 | 0.39 | 0.38 | 2–4 | 41.2 | 0.35 | –1.17 | 0.0524 | 2.690 | [8] |
Sheepshead | 20 | 0.23 | 0.22 | 2–5 | 41.9 (M) 44.7 (F) |
0.417 0.367 |
–0.901 to 1.025 |
0.000448 0.000530 |
2.88 2.85 |
[9] |
Atlantic spadefish |
8 | 0.41 | 0.58 | 1–8 | 49.0 | 0.340 | –0.18 | 0.0927 | 2.64 | [10] |
Bluefish | 8 | 0.41 | 0.58 | 1–6 | 94.4 | 0.18 | 1.033 | –10.02 | 2.80 | [11] |
where
where
where G is the specific growth rate in kg yr-1, W is weight in kg and t is time.
Species | Trophic Level (SE) |
Diet Composition (source) | IRE | T°C | Pr | Pe | APr | APe |
---|---|---|---|---|---|---|---|---|
Red snapper | 4.01 ± 0.59 | Benthic inverts, demersal fishes, squid, pelagic zooplankton [12–15] |
0.05 | 25.9 | 306 | 115 | 15 | 6 |
Blue runner | 4.40 ± 0.77 | Fish, decapods, hyperid amphipods, chaetognaths, other [[16], FishBase] |
0.10 | 25.9 | 1627 | 333 | 163 | 33 |
Sheepshead | 3.53 ± 0.53 | Portunid crabs, shrimp, barnacles, fish, copepods, bryozoans, amphipods, sargassum [[17], FishBase] |
0.90 | 25.9 | 410 | 185 | 369 | 167 |
Atlantic spadefish |
3.50 ± 0.47 | Sponges and tunicates, cnidarians, worms, ascidicans, plants, benthic inverts, echinoderms, zooplankton (FishBase) |
0.95 | 25.9 | 987 | 243 | 938 | 231 |
Bluefish | 4.50 ± 0.55 | Pelagic and demersal fish and macrocrustaceans (from soft bottoms) [18] |
0.01 | 25.9 | 561 | 164 | 6 | 2 |
3. Results
The species for which we assigned an N have life history and behavioral characteristics that are qualitatively similar to the attraction end of the Bohnsack’s continuum [1]. These species are directly fished or overfished, exhibit low site fidelity, are less or not dependent upon the reef for food, are not dependent upon the reef for completion of their life cycles, and are pelagic and/or migratory, and thus less likely to habitat limited. In contrast, the species for which we assigned a Y in ATable 1 have life history and behavioral characteristics that are qualitatively more similar to production end of the continuum [1]. These species have relatively high site fidelity, the need for reef or structured habitat to complete the life cycle, and a significant fraction of their diet is comprised primarily of reef-associated prey.
There are numerous species for which expectations about reef dependence are more difficult to describe, even qualitatively. To provide some interpretation, we use both a qualitative assessment relative to the Bohnsack [1] conceptual model, and insight derived from the Level-3 assessments to make comparisons among the reef-associated species in ATable 1. Where possible, we identify species that are comparable with respect to ecology, life history, and behavioral characteristics to the Level-3 species. It is fortunate that the latter group is comprised of species that appear to differ significantly in their relative ecological dependence on reefs and, by extension, to platforms. We also consulted additional reference materials to make our determinations [18–24].
In all, 46 species are listed as reef associated (RA in ATable 1). Of these, many are known to be pelagic and/or highly migratory. Among this group are several species of jacks (fm. Carangidae, genus
Thirty of the 46 species listed as reef associated (RA) have other habitats listed as primary (fm. Carangidae, mackerels fm. Scombribdae, and clupeiods herrings and anchovies). Many of these species are reported to primarily associate with hard-bottom (HB) habitats (25 species including most of the groupers). This makes sense given the nature of most of the natural reef habitat in the GOM. Other species are reported to associate with sea grass (SG) meadows. Where possible, we have included additional detail in ATable 1 about primary habitat associations reported for many of the reef-associated species. These primary habitats are consistent with natural habitats reported to occur in the GOM and include reef flats, rocky reefs, coral reefs, oyster reefs, floatsam, shelf-edge banks, offshore rock bottoms, offshore banks,
Most of the grouper species reported in ATable 1 in the genus
Gag groupers are more widely distributed in the GOM, but also are overfished [28]. They are extremely vulnerable to overexploitation because they are haremic as adults, and aggregate to spawn at just a few locations in the northeastern GOM. Juvenile gag groupers are mostly associated with sea grass meadows as nursery areas. To our knowledge, the ecology of gag grouper on platforms has received little study. However, the work of Lindberg and coworkers on the west Florida shelf has demonstrated that the value of artificial reefs as habitat is affected both by size and spatial arrangement of reef modules. The net effect on stock production of reefs is negative when fishing mortality is considered [29, 30]. Despite these results, we caution against drawing inference about the role of platforms as habitat for gag groupers because the aforementioned work was done on relatively small, low-relief, reef modules. Two other species reported as reef-associated in ATable 1 (
There are several species in ATable 1 that are reported to be reef-associated, but also occur on a wide variety of habitats including inshore waters, bays, estuaries, and sea grass meadows. Qualitatively, these species have life history and behavioral characteristics that are more similar to those found at the production end of Bohnsack’s continuum [1]. These include
Similarly, there is another group that qualitatively appears to have life history and behavioral characteristics that are more similar to those found at the production end of the continuum, but also appear to be more restricted in their distribution than the reef-associated group that use many habitats (discussed in the previous paragraph). These more habitat-restricted species are reported to occur in coastal waters, and on shelf-edge banks, but are explicitly identified as not being found on coral reefs. This group includes two members of the genus
There are several small, cryptic species listed in ATable 1 as reef-associated that we believe to be more strongly associated with reefs than the many-habitat and restricted habitat reef-associated species (preceding two paragraphs), and whose life history and behavioral characteristics appear to place them solidly at the production end of Bohnsack’s continuum. This group includes
In addition, there are several species listed in ATable 1 as being reef-associated in FishBase, but whose life history and behavioral characteristics (at least qualitatively) do not strongly support placement near either endpoint of Bohnsack’s continuum. This group has been reported to occur on a wide variety of natural hard-bottom habitats in the GOM, including platforms, but appear to have only moderate to low site fidelity. Many of these species support directed commercial fisheries, and all appear among the list of species harvested by recreational anglers. This group includes six members of the genus
Of this group only the red snapper, and to some degree
The other lutjanids in this group are much less abundant than red and vermilion snapper. Vermilion snapper,
Calculated estimates of biomass production per year per platform using Rickers’s method ranged from 306 kg platform-1 by sheepshead to 1627 kg platform-1 for blue runner (Table 3). Estimates using an empirical approach [3] were consistent in pattern, but averaged less than half of the values derived from Ricker’s [4] method (Table 3). This difference was largely because we estimated specific growth rates (G) for each species over a period that was shorter than their reported life span. For red snapper, blue runner, and sheepshead, the age classes we used were those in which high growth occurred during that period of their life cycle. Biomass production in the Ricker’s method is sensitive to the ratio of G/Z i.e., specific growth rate and natural mortality rate, and the ratio is close to, or less than, one for all, but red snapper (Table 3). When the ratio is less than one, there is a net loss in population biomass. It is also important to note that the often highly productive pre-recruit period was not included in our calculations, which could have a large effect on the overall production estimates.
The results from the Powers [2] model showed that annual production for each species was dependent upon the index of reef exclusivity (Table 3). Species for which platforms provide only a small fraction of prey resources (e.g., red snapper, blue fish, and blue runner) are less dependent upon reefs compared to species such as Atlantic spadefish and sheepshead that depend heavily upon the fouling community on platform legs for food. Low annual production values for red snapper, bluefish, and blue runner also imply that platforms are more likely attracting individuals from surrounding natural habitats rather than producing new population biomass. These finding are consistent with recent diet studies of red snapper, and point out the sensitivity of production estimates to subtle changes in the G/Z ratio that are not considered in the less complex methods [2, 3].
4. Final Thoughts and Red Snapper
The most controversial fishery in U.S. waters of the GOM is for northern red snapper
Like the examples described by Hilborn [40], a ‘faith-based fisheries’ argument has been used to defer effective management of red snapper, and consequently has greatly strained the relationships among science, management, and stakeholders in the GOM. It has been argued that mass deployment of artificial reefs has substantially increased productivity of the red snapper stock. The premise is that artificial reefs have transformed less desirable fish biomass into red snapper biomass at locations on the shelf where the latter was not previously abundant [41]. The specifics of this argument were elucidated in a management perspective [42] which postulated that oil and gas platforms that began appearing in the western GOM in the late 1940s function secondarily as large artificial reefs, as well as a myriad of other artificial reef structures in the northcentral and eastern GOM since the 1970s, has enhanced biomass production of red snapper. True, results of the last benchmark stock assessment for red snapper [7] indicate that recruitment and stock productivity may have increased since the late 1980s. However, several other possible causes for this putative increase have been identified beyond artificial reef deployment [39]. The perspective [42] speaks to none of the other possible (more likely?) causes for change and further claims that red snapper were not present in the northwestern GOM until oil and gas platforms began being deployed offshore in the early 1940s. They disregard considerable information showing that a well-established red snapper fishery in the northwestern GOM began as early as 1892 [43].
The arrows in Figure 3 indicate when artificial reefs began to be deployed in large numbers relative to the estimated spawning stock biomass of red snapper. These deployments took the form of oil and gas platforms in the western GOM and all manner of materials in the east. In both cases, there is no obvious indication that artificial habitats have increased spawning stock biomass because overfishing was occurring until only recently, and changed in response to strong year classes. The artificial reef argument put forth in the perspective is simply not supported by the available information.
We recognize that the stock assessment process for red snapper and other reef-associated species is controversial and sometimes difficult to understand, so in this chapter we have used simple models using a few parameters, to make our case. In Table 3 our results indicate that oil and gas platforms produce new red snapper biomass. However, even if only small increase in Z in the Ricker based method of estimation [4] is added (i.e., as fishing mortality (Z=F + M)), the production outcome for red snapper turns negative. We provided tables that include procedural details for readers that wish to see how we estimated production of red snapper on oil and gas platforms. It is entirely plausible that oil and gas platforms make some reef-associated species, including red snapper, more vulnerable to fishing. Our analysis does not address spatial variation in demographic rates due to the intrinsic habitat quality of essential fish habitat or artificial reefs at scales smaller than regional e.g., between Alabama, Louisiana, and Texas.
This is not a blanket recrimination of artificial reefs and artificial reef programs. There are clear examples in the literature where artificial reefs benefit fishes and ecosystems in which they have been employed. Good examples are where artificial reefs are used to mitigate for loss or injury to natural reefs, or used to reduce destructive diving and fishing pressure on natural reefs [44–47], to name a few.
Still, the debate about whether artificial habitats attract red snapper from nearby natural habitats or actually enhance production of new biomass (i.e., the attraction vs. production debate) has been called meaningless and unresolvable [40]. This subject often is debated in broad form for all reef-associated species and, as such, may be un-resolvable in the broader context—this issue has mostly been tried in the court of public opinion. However, a more quantitative approach is tractable for a well studies species like red snapper, although difficult due to the scale and complexity of needed studies [48].
Besides red snapper, many reef-associated species that are found on oil and gas platforms and highlighted in red in ATable 1 are overfished. The role that these structures play in the population dynamics of these species is unknown. Careful consideration and enumeration of potential positive and negative impacts of man-made habitats on dynamic coastal geographies on continental shelves should be made before such habitats are constructed.
Appendices
Taxa | SF | DF | Diet | PC | Behavior | Dep. | |
---|---|---|---|---|---|---|---|
H | N | R | R? | R, D | Y | ||
H | N | R | R? | R, D | Y | ||
L | Y | P | R | P, HM | N | ||
H | N | R | R | R, D | Y | ||
H | N | R | R? | R, D | Y | ||
L | N | B | R | D | N | ||
L | Y | B | R | D | N | ||
H | N | R, B | R? | R, D | Y | ||
H | N | R, B | R? | R, D | Y | ||
H | N | R, B | R? | R, D | Y | ||
L | N | P | R | P | N | ||
L | N | P | R | P | N | ||
L | N | P | R | P | N | ||
L | N | P | R | P | N | ||
L | N | P | R | P | N | ||
L | N | B | R | D | N | ||
L | Y | B | R | D | N | ||
H | N | R, B | H | R, D | Y | ||
H | N | R, B | R? | R, D | Y | ||
H | N | R, B | R? | R, | Y | ||
M | Y | R, B | R | R, D | RA | ||
L | N | B | R | P | S | ||
L | N | B | R | D | N | ||
L | Y | B, P | R | D | N | ||
L | N | B, P | R | D | N | ||
H | Y | R, B | H | R, D | Y | ||
H | Y | R, P | R? | R, P | Y | ||
L | N | B | R | D | N | ||
L | N | B | R | D | N | ||
L | Y | P | R | P | N | ||
L | N | B | R | D | RA (juv) |
||
M? | N | B? | R? | D | RA, SG | ||
H | N | B | H? | R, D (adults, juv=P) |
Y (adults) |
||
L | N | P | R | D, P | N, SG, RA | ||
H? | N | B | H? | R, D | Y?, RA & SG | ||
L | N | P | R | HM | RA | ||
L | N | P | R | P | RA | ||
L | N | P | R | P | RA | ||
L | N | P | R | P | RA | ||
L | N | P | R | P | RA | ||
L | N | P | R | P | RA | ||
L | Y | P | R | P, HM | N | ||
L | Y | P, B | R | D | N | ||
M? | Y | P, B | R | D | RA | ||
M? | N | B | R | R, D | HB | ||
M? | N | B | R | P, D | HB | ||
H | N | P | H? | R, D | Y | ||
H | N | B, P | R | P, P | Y? | ||
H | N | R | H? | R, D | Y | ||
H | N | R | H? | R, D | Y | ||
L | N | P | R | P | N | ||
L | N | P | R | P | N | ||
M | N | B | R | D | RA, SG | ||
L | N | B | R | D | N | ||
L | N | P | R | P | N | ||
H | N | P | H | D | Y | ||
H | N | R | H | D | Y | ||
H | N | R | H | D | Y | ||
L | Y | B | R | D | N | ||
H | N | P | H | D, P | Y | ||
L | Y | P | R | P, HM | N | ||
L | Y | P | R | P, HM | N | ||
M | N | ? | R | D | RA | ||
L | N | P | R | P | N | ||
L | Y | B | R | D | N | ||
L | Y | ? | R | D | N | ||
L | N | P | R | P | N | ||
L | Y | P | R | P | N | ||
L | Y | P | R | P | N, RA | ||
L | Y | P | R | P | N, RA | ||
L | Y | P | R | P, D | N | ||
H? | N | R? | H | D | Y, HB | ||
H | Y?* | R, P | H? | D | RA, HB | ||
L | N | B | R | D | N | ||
L | N | P | R | D | SG, HB | ||
H | N | B | R | D | SG | ||
L | N | P | R | P | N | ||
L | N | B | R | D | N, SG | ||
L | N | P (omnivore) |
R | D | N | ||
L | N | P | R | P | N, RA | ||
L | N | P | R | P | N | ||
L | N | B | R | D | N | ||
M | Y | P | R | P, HM | RA | ||
L | N | P | R | P | N | ||
L | N | P | R | P | N | ||
L? | N? | ? | R | D | N? | ||
H | Y?* | R, P, B | H | D | Y, HB | ||
M | Y | R, P, B | R | D | RA | ||
M | Y | R, B | R | D | RA, HB | ||
M | N | R, P, B | R | D | RA, HP | ||
(closed) | |||||||
M | Y* | P, B | R | D | RA, HB | ||
M? | N | B | R? | D | RA? | ||
M | N | B | R? | D | RA? | ||
L | N | B | R | D | N | ||
L | N | P | R | P | N | ||
L | N | P | R | P, HM | N, RA | ||
L | N | B | R | D | N | ||
M? | N | B | R | D | N, SG | ||
(sometimes found on coral reefs) | |||||||
M? | N | B | R | D | N, SG, HB | ||
(common on oyster reefs) | |||||||
L | N | B? | R | D | N | ||
H | N | P, B | R? | D | |||
H | N | P, B | R | D | Y?, SG, HB | ||
M | Y | P, B | R | D | |||
H | N | P, B | H? | D | Y, HB | ||
L | N | B? | R | D | N | ||
M | N | P, B | R | P | RA | ||
H | N | B | R? | D | Y | ||
H | N | B | R? | D | Y | ||
H | N | B | R? | D | Y, RA | ||
H | N | B | R? | D | Y | ||
L? | N | B? | R? | D | N?, RA | ||
L | N | P | R | P | N | ||
H | N | B? | H | D | RA, HB | ||
(nest-builder) | |||||||
H | N | B? | H | D | RA, HB | ||
(nest-builder) | |||||||
H | N | B? | H | D | RA, HB | ||
(nest-builder) | (common on oyster reefs) | ||||||
H | N | B | H | D | RA, HB | ||
(nest-builder) | |||||||
H | N | B? | H | D | RA, HB | ||
(nest-builder) | |||||||
L | N | P | R | D | N | ||
L | Y | P | R | P | N? | ||
(associated with objects drifting at surface) | |||||||
M? | N | P? | R | P | Y?, HB. S | ||
(plants, including Sargassum) | |||||||
M? | N | P, B | R | P | Y?, HB, S | ||
(plants, including Sargassum, some benthic crustaceans) | |||||||
M? | Y | B | R | D | Y?, HB | ||
L | N | B | R | D | SG | ||
M | N | B | R | D | RA, SG, HB | ||
L | N | B | R | D | N | ||
L | Y | B | R | D | N | ||
L | N | B | R | D | N | ||
L | N | ? | R | D | N | ||
(deep water) | |||||||
L | N | ? | R | P | N | ||
(bathypelagic) | |||||||
M | Y | P, B | R | P | RA | ||
(often found associated with flotsam) | |||||||
M | Y | P, B | R | D | RA | ||
M | Y | P, B | R | D, P | RA, HB | ||
M | Y | P | R | P | RA | ||
M | Y | R, P, B | R | P | RA, HB | ||
M | Y | R, P, B | R | D | RA, HB | ||
M | Y | R, P, B | R | D | RA, HB | ||
(common on shelf-edge banks) | |||||||
L | N | P | R | P | N | ||
(bathypelagic) | |||||||
L | Y | P | R | P, HM | N | ||
L | Y | P | R | P | N, RA | ||
L | N | P | R | P | N | ||
L | N | B? | R | D | N? | ||
L | N | B? | R | D | N | ||
L | Y | B | R | D | N | ||
L | N | B | R | D | N | ||
(bathydemersal) | |||||||
L | Y | B | R | P | N | ||
(plants) | |||||||
L | Y | B | R | P | N | ||
L | N | B | R | D | N | ||
M | Y | P | R | P, D | RA, SG, HB | ||
adults offshore on rocky bottoms) | |||||||
M | Y* | R, P | R | D | RA, HB | ||
(high-relief rocky bottoms, often found on Oculina reefs) | |||||||
M | Y* | R, P, B | R | D | RA, HB | ||
(rocky and sandy bottoms) | |||||||
M | Y* | R, P | R | P, D | RA | ||
(rocky and coral reefs, shelf-edge banks in GOM) | |||||||
L | N | B? | R | D | RA, SG | ||
M | N | B | R | D | RA (offshore banks) | ||
M | Y | R, P, B | R | P | RA (mostly coral reefs) | ||
L | N | B? | R | D | N, HB | ||
L | N | B? | R | D | N, HB | ||
L | N | P, B | R | P | RA | ||
L | N | B | R | D | N | ||
(common on shrimp grounds) | |||||||
L | N | B? | R | D | N | ||
(uncommon, shallow sandy bays) | |||||||
L? | N | B? | R? | D | N | ||
M? | N | B? | R? | D | RA | ||
H | N | B | H | D | RA | ||
(plants) | (rocky reefs and corals) | ||||||
H | N | R, P | R? | D | Y? | ||
H | N | R? | R? | D | Y? | ||
M | N | B | H? | D | RA, SG | ||
` | (common on oyster reefs) | ||||||
M | N | B | H? | D | RA, HB | ||
L | N | P, B | R | D | N | ||
H | N | B | H | D | Y | ||
(mostly algae) | |||||||
L | Y | P, B | R | D | N, HB | ||
L | Y | P, B | R | D | N | ||
H | N | P | R | P | RA, HB | ||
(coral reefs, hard bottoms) | |||||||
L | N | B | R | D | N | ||
L | N | P | R | P | N | ||
L | N | P? | R | P | N | ||
(bathypelagic) | |||||||
L | Y | P | R | P, D | N | ||
(benthopelagic) | |||||||
L | Y | B | R | D | N | ||
L | N | B | R | D | N | ||
H | N | R, B | R? | D | Y | ||
L | Y | P | R | P | N | ||
L | N | B | R | D | N | ||
H | N | R, P, B | R? | D | Y | ||
L | N | B | R | D | N | ||
L | N | P | R | D | N | ||
(small fishes) | |||||||
H | N | R? | H | D | HB | ||
(offshore banks) | |||||||
M? | N | B | R? | D | RA, HB | ||
M | Y | P, B | R | P | RA | ||
(found associated with structure of all types) | |||||||
L | N | B | R | D | N | ||
H? | N | P | R | P | RA? | ||
(usually attached to sharks, turtles) | |||||||
M | Y | P, B | R | P | RA, HB | ||
(HB on shelf-edge) | |||||||
L | N | P | R | P | N | ||
(bathypelagic) | |||||||
L | N | P, B | R | P | N | ||
(bathypelagic) | |||||||
M? | N | B | R | D | RA? | ||
L | Y | P | R | P | RA | ||
L | N | P | R | D | N | ||
(nekton) | |||||||
M | N | P | R | D | RA | ||
L | N | P | R | ||||
H | N | R, B | H | D | Y | ||
L | N | P | R | P | N | ||
L | Y | P, B | R | D, P | N | ||
L | Y | P | R | P, HM | N | ||
L | Y | P | R | P, HM | RA | ||
Y | P | R | P, HM | RA | |||
L | N | B | R | D | HB | ||
L | N | P | R | P | RA | ||
L | N | P, B | R | P | N | ||
(benthopelagic) | |||||||
L | N | P, B | R | P | N | ||
M | Y | P | R | P, HM | RA | ||
M | Y | P | R | P, HM | RA | ||
M | Y | P | R | P, HM | RA | ||
M | Y | P | R | P, HM | N | ||
L | N | B | R | D | N | ||
L | N | B | R | D | N | ||
L | N | B | R | D | RA | ||
(SG, reef flats) | |||||||
M | Y | P | R | P | RA, SG | ||
M | N | P | R | P | RA | ||
L | N | P | R | P | N | ||
H | N | R, B | R? | D | Y | ||
H | N | R, B | R? | D | Y | ||
L | N | B | R | D | N | ||
L | N | B | R | D | N | ||
H? | N | B | R? | P, D | RA | ||
M | N | B, P | R | D | RA | ||
M | N | B | R | D | RA | ||
L | N | B? | R | D | N | ||
H | N | P | R | D | RA | ||
L | N | B | R | D | HB | ||
L | N | B | R | D | RA, HB | ||
M | N | P | R | D | RA, HB | ||
(nekton) | |||||||
L | N | P | R | P | N | ||
(jellyfish) | |||||||
H | N | R, B | R | D | RA, SG | ||
L | Y | P | R | P, HM | N, RA | ||
L | Y | P | R | P, HM | N, RA | ||
L | Y | P | R | P, HM | N | ||
M | N | B, P | R | D | RA, HB | ||
L | Y | B, P | R | D | N | ||
(benthopelagic) | |||||||
M | Y | B | R | D | RA | ||
M | Y | B, P | R | P | RA | ||
(benthopelagic) | |||||||
M | Y | P | R | P | RA | ||
(minor, bait) | |||||||
L | Y | P, B | R | P, D | N | ||
L | Y | B | R | D | N | ||
L | N | B | R | D | N | ||
L | N | P, B | R | D | N | ||
(zoobenthos, small fishes) | |||||||
L | N | B, P | R | D | N | ||
L | N | P | R | P | N | ||
(bathypelagic) | |||||||
H | N | B | H? | D | Y? |
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