Biogeographic characterization of fish communities and associated benthic habitats within the Flower Garden Banks National Marine Sanctuary: sampling design and implementation of scuba surveys on the coral caps

The Flower Garden Banks National Marine Sanctuary (FGBNMS) is located in the northwestern Gulf of Mexico approximately 180 km south of Galveston, Texas. The sanctuary’s distance from shore combined with its depth (the coral caps reach to within approximately 17 m of the surface) result in limited exposure of this coral reef ecosystem to natural and human-induced impacts compared to other coral reefs of the western Atlantic. In spite of this, the sanctuary still confronts serious impacts including hurricanes events, recent outbreaks of coral disease, an increase in the frequency of coral bleaching and the massive Diadema antillarum die-off during the mid-1980s. Anthropogenic impacts include large vessel anchoring, commercial and recreational fishing, recreational scuba diving, and oil and gas related activities. The FGBNMS was designated in 1992 to help protect against some of these impacts. Basic monitoring and research efforts have been conducted on the banks since the 1970s. Early on, these efforts focused primarily on describing the benthic communities (corals, sponges) and providing qualitative characterizations of the fish community. Subsequently, more quantitative work has been conducted; however, it has been limited in spatial scope. To complement these efforts, the current study addresses the following two goals put forth by sanctuary management: 1) to develop a sampling design for monitoring benthic fish communities across the coral caps; and 2) to obtain a spatial and quantitative characterization of those communities and their associated habitats.

A point for comparison: the Flower Garden Banks National Marine Sanctuary

A reoccurring goal listed during the creation of Marine Protected Areas (MPAs) is to return the region to a former state. However, limited data is available that describes or characterizes this former condition. Data collected from ecosystems with comparatively limited anthropogenic impacts, can provide invaluable information in suggesting what former states may have looked like. One example is the Flower Garden Banks National Marine Sanctuary which is located 180 kilometers off the coast of Texas. These relatively isolated and pristine banks are capped by substantial scleractinian coral communities, forming excellent habitat for over 200 species of fish. While fishing is permitted, it is limited by difficulty of access. In 2006, NOAA’s Biogeography Branch, in collaboration with the Sanctuary, initiated the first quantitative assessment of fish resources throughout the diveable portions of the Sanctuary. The sampling design and methodologies employed were identical to those that the Branch has utilized in other more impacted regions of the US Caribbean. Initial analyses reveal that fish density and species richness at the Sanctuary were almost two times greater than that found within the US Caribbean and biomass was approximately six times higher. This was due in large part to the presence of sizeable piscivores of the genera Mycteroperca and Dermatolepis. The Sanctuary is one of few minimally impacted locations remaining within the Tropical Western Atlantic. As such, these findings should be considered when attempting to establish a former state or evaluate effectiveness of an MPA in meeting its management goals.

Status of the coral reef ecosystems in the U.S. Caribbean and Gulf of Mexico: Florida, Flower Garden Banks, Puerto Rico, Navassa and USVI

This chapter covers coral reef areas under the jurisdiction of the USA in the Wider Caribbean: Florida; Flower Garden Banks; Puerto Rico; U.S. Virgin Islands; and Navassa. The following information is condensed from six chapters of The State of Coral Reef Ecosystems of the United States and Pacific Freely Associated States: 2008. Access to the full text of this comprehensive report is available at: http://ccma.nos.noaa.gov/stateofthereefs.

Historical nitrogen and phosphorus loadings to the northern Gulf of Mexico

Primary productivity in many coastal systems is nitrogen (N) limited; although, phytoplankton productivity may be limited by phosphorus (P) seasonally or in portions of an estuary. Increases in loading of limiting nutrients to coastal ecosystems may lead to eutrophication (Nixon 1996). Anthropogenically enhanced eutrophication includes symptoms such as loss of seagrass beds, changes in algal community composition, increased algal (phytoplankton) blooms (Richardson et al. 2001), hypoxic or anoxic events, and fish kills (Bricker et al. 2003).

Spatial and temporal variation in the diet of the California sea lion (Zalophus californianus) in the Gulf of California, Mexico

Between June 1995 and May 1996 seven rookeries in the Gulf of California were visited four times in order to collect scat samples for studying spatial and seasonal variability California sea lion prey. The rookeries studied were San Pedro Mártir, San Esteban, El Rasito, Los Machos, Los Cantiles, Isla Granito, and Isla Lobos. The 1273 scat samples collected yielded 4995 otoliths (95.3%) and 247 (4.7%) cephalopod beaks. Fish were found in 97.4% of scat samples collected, cephalopods in 11.2%, and crustaceans in 12.7%. We identified 92 prey taxa to the species level, 11 to genus level, and 10 to family level, of which the most important were Pacific cutlassfish (Trichiurus lepturus), Pacific sardine (Sardinops caeruleus), plainfin midshipman (Porichthys spp.), myctophid no. 1, northern anchovy (Engraulis mordax), Pacific mackerel (Scomber japonicus), anchoveta (Cetengraulis mysticetus), and jack mackerel (Trachurus symmetricus). Significant differences were found among rookeries in the occurrence of all main prey (P≤0.04), except for myctophid no. 1 (P>0.05). Temporally, significant differences were found in the occurrence of Pacific cutlassfish, Pacific sardine, plainfin midshipman, northern anchovy, and Pacific mackerel (P<0.05), but not in jack mackerel (χ 2=2.94, df=3, P=0.40), myctophid no. 1 (χ 2=1.67, df= 3, P=0.64), or lanternfishes (χ 2=2.08, df=3, P=0.56). Differences were observed in the diet and in trophic diversity among seasons and rookeries. More evident was the variation in diet in relation to availability of Pacific sardine.

Growth, mortality, and hatchdate distributions of larval and juvenile spotted seatrout (Cynoscion nebulosus) in Florida Bay, Everglades National Park

Life history aspects of larval and, mainly, juvenile spotted seatrout (Cynoscion nebulosus) were studied in Florida Bay, Everglades National Park, Florida. Collections were made in 1994−97, although the majority of juveniles were collected in 1995. The main objective was to obtain life history data to eventually develop a spatially explicit model and provide baseline data to understand how Everglades restoration plans (i.e. increased freshwater flows) could influence spotted seatrout vital rates. Growth of larvae and juveniles (<80 mm SL) was best described by the equation loge standard length = –1.31 + 1.2162 (loge age). Growth in length of juveniles (12–80 mm SL) was best described by the equation standard length = –7.50 + 0.8417 (age). Growth in wet weight of juveniles (15–69 mm SL) was best described by the equation loge wet-weight = –4.44 + 0.0748 (age). There were no significant differences in juvenile growth in length of spotted seatrout in 1995 between three geographical subdivisions of Florida Bay: central, western, and waters adjacent to the Gulf of Mexico. We found a significant difference in wet-weight for one of six cohorts categorized by month of hatchdate in 1995, and a significant difference in length for another cohort. Juveniles (i.e. survivors) used to calculate weekly hatchdate distributions during 1995 had estimated spawning times that were cyclical and protracted, and there was no correlation between spawning and moon phase. Temperature influenced otolith increment widths during certain growth periods in 1995. There was no evidence of a relationship between otolith growth rate and temperature for the first 21 increments. For increments 22–60, otolith growth rates decreased with increasing age and the extent of the decrease depended strongly in a quadratic fashion on the temperature to which the fish was exposed. For temperatures at the lower and higher range, increment growth rates were highest. We suggest that this quadratic relationship might be influenced by an environmental factor other than temperature. There was insufficient information to obtain reliable inferences on the relationship of increment growth rate to salinity.

A remarkable new species of Psednos (Teleostei: Liparidae) from the western North Atlantic Ocean

Psednos rossi new species (Teleostei: Liparidae) is described from two specimens collected in the North Atlantic Ocean off Cape Hatteras, North Carolina, at depths of 500–674 m. Psednos rossi belongs to the P. christinae group, which includes six other species and is characterized by 46–47 vertebrae and the absence of a coronal pore. Psednos rossi differs from those six species by having characters unique within the genus: straight spine, body not humpbacked at the occiput, and a very large mouth with a vertical oral cleft. Other distinguishing characters include a notched pectoral fin with 15–16 rays, eye 17–19% SL, and color in life orange-rose. With P. rossi, the genus Psednos as currently known includes 26 described and five undescribed species of small meso- or bathypelagic liparids from the Atlantic, Pacific, and Indian Oceans.

Sizes of walleye pollock (Theragra chalcogramma) and Atka mackerel (Pleurogrammus monopterygius) consumed by the western stock of Steller sea lions (Eumetopias jubatus) in Alaska from 1999 to 2000

Prey-size selectivity by Steller sea lions (Eumetopias jubatus) is relevant for understanding the foraging behavior of this declining predator, but studies have been problematic because of the absence and erosion of otoliths usually used to estimate fish length. Therefore, we developed regression formulae to estimate fish length from seven diagnostic cranial structures of walleye pollock (Theragra chalcogramma) and Atka mackerel (Pleurogrammus monopterygius). For both species, all structure measurements were related with fork length of prey (r2 range: 0.78−0.99). Fork length (FL) of walleye pollock and Atka mackerel consumed by Steller sea lions was estimated by applying these regression models to cranial structures recovered from scats (feces) collected between 1998 and 2000 across the range of the Alaskan western stock of Steller sea lions. Experimentally derived digestion correction factors were applied to take into account loss of size due to digestion. Fork lengths of walleye pollock consumed by Steller sea lions ranged from 3.7 to 70.8 cm (mean=39.3 cm, SD=14.3 cm, n=666) and Atka mackerel ranged from 15.3 to 49.6 cm (mean=32.3 cm, SD=5.9 cm, n=1685). Although sample sizes were limited, a greater proportion of juvenile (≤20 cm) walleye pollock were found in samples collected during the summer (June−September) on haul-out sites (64% juveniles, n=11 scats) than on summer rookeries (9% juveniles, n=132 scats) or winter (February−March) haul-out sites (3% juveniles, n=69 scats). Annual changes in the size of Atka mackerel consumed by Steller sea lions corresponded to changes in the length distribution of Atka mackerel resulting from exceptionally strong year classes. Considerable overlap (>51%) in the size of walleye pollock and Atka mackerel taken by Steller sea lions and the sizes of these species caught by the commercial trawl fishery were demonstrated.

Red snapper (Lutjanus campechanus) demographic structure in the northern Gulf of Mexico based on spatial patterns in growth rates and morphometrics

Red snapper (Lutjanus campechanus) in the United States waters of the Gulf of Mexico (GOM) has been considered a single unit stock since management of the species began in 1991. The validity of this assumption is essential to management decisions because measures of growth can differ for nonmixing populations. We examined growth rates, size-at-age, and length and weight information of red snapper collected from the recreational harvests of Alabama (n=2010), Louisiana (n=1905), and Texas (n =1277) from 1999 to 2001. Ages were obtained from 5035 otolith sections and ranged from one to 45 years. Fork length, total weight, and age-frequency distributions differed significantly among all states; Texas, however, had a much higher proportion of smaller, younger fish. All red snapper showed rapid growth until about age 10 years, after which growth slowed considerably. Von Bertalanffy growth models of both mean fork length and mean total weight-at-age predicted significantly smaller fish at age from Texas, whereas no differences were found between Alabama and Louisiana models. Texas red snapper were also shown to differ significantly from both Alabama and Louisiana red snapper in regressions of mean weight at age. Demographic variation in growth rates may indicate the existence of separate management units of red snapper in the GOM. Our data indicate that the red snapper inhabiting the waters off Texas are reaching smaller maximum sizes at a faster rate and have a consistently smaller total weight at age than those collected from Louisiana and Alabama waters. Whether these differences are environmentally induced or are the result of genetic divergence remains to be determined, but they should be considered for future management regulations.

Descriptions of larval, prejuvenile, and juvenile finescale menhaden (Brevoortia gunteri) (family Clupeidae), and comparisons to gulf menhaden (B. patronus)

Larval and juvenile development of finescale menhaden (Brevoortia gunteri) is described for the first time by using wild-caught individuals from Nueces Bay, Texas, and is compared with larval and juvenile development of co-occurring gulf menhaden (B. patronus). Meristics, morphometrics, and pigmentation patterns were examined as development proceeded. An illustrated series of finescale menhaden is presented to show changes that occurred during development. For finescale menhaden, transformation to the juvenile stage was completed by 17−19 mm standard length (SL). By contrast, transformation to the juvenile stage for gulf menhaden was not complete until 23−25 mm SL. Characteristics useful for separating larval and juvenile finescale menhaden from gulf menhaden included 1) the presence or absence of pigment at the base of the insertion of the pelvic fins; 2) the standard length at which medial predorsal pigment occurs; 3) differences in the number of dorsal fin ray elements; and, 4) the number of vertebrae.

Small-scale spatial and temporal variability in growth and mortality of fish larvae in the subtropical northcentral Gulf of Mexico: implications for assessing recruitment success

Extensive plankton collections were taken during seven September cruises (1990–93) along the inner continental shelf of the northcentral Gulf of Mexico (GOM). Despite the high productivity and availability of food during these cruises, significant small-scale spatial variability was found in larval growth rates for both Atlantic bumper (Chloroscombrus chrysurus, Carangidae) and vermilion snapper (Rhomboplites aurorubens, Lutjanidae). The observed variability in larval growth rates was not correlated with changes in water temperature or associated with conspicuous hydrographic features and suggested the existence of less-recognizable regions where conditions for growth vary. Cruise estimates of mortality coefficients (Z) for larval Atlantic bumper (n=32,241 larvae from six cruises) and vermilion snapper (n= 2581 larvae from four cruises) ranged from 0.20 to 0.37 and 0.19 to 0.29, respectively. Even in a subtropical climate like the GOM, where larval-stage durations may be as short as two weeks, observed variability in growth rates, particularly when combined with small changes in mortality rates, can cause order-of-magnitude differences in cumulative larval survival. To what extent the observed differences in growth rates at small spatial scales are fine-scale “noise” that ultimately is smoothed by larger-scale processes is not known. Future research is needed to further characterize the small-scale variability in growth rates of larvae, particularly with regard to microzooplankton patchiness and the temporal and spatial pattern of potential predators. Small-scale spatial variability in larval growth rates may in fact be the norm, and understanding the implications of this subtle mosaic may help us to better evaluate our ability to partition the causes of recruitment variability.

Prey consumption of Steller sea lions (Eumetopias jubatus) off Alaska: How much prey do they require?

The effects of seasonal and regional differences in diet composition on the food requirements of Steller sea lions (Eumetopias jubatus) were estimated by using a bioenergetic model. The model considered differences in the energy density of the prey, and differences in digestive efficiency and the heat increment of feeding of different diets. The model predicted that Steller sea lions in southeast Alaska required 45–60% more food per day in early spring (March) than after the breeding season in late summer (August) because of seasonal changes in the energy density of the diets (along with seasonal changes in energy requirements). The southeast Alaska population, at 23,000 (±1660 SD) animals (all ages), consumed an estimated 140,000 (±27,800) t of prey in 1998. In contrast, we estimated that the 51,000 (±3680) animals making up the western Alaska population in the Gulf of Alaska and Aleutian Islands consumed just over twice this amount (303,000 [±57,500] t). In terms of biomass removed in 1998 from Alaskan waters, we estimated that Steller sea lions accounted for about 5% of the natural mortality of gadids (pollock and cod) and up to 75% of the natural mortality of hexagrammids (adult Atka mackerel). These two groups of species were consumed in higher amounts than any other. The predicted average daily food requirement per individual ranged from 16 (±2.8) to 20 (±3.6) kg (all ages combined). Per capita food requirements differed by as much as 24% between regions of Alaska depending on the relative amounts of low–energy-density prey (e.g. gadids) versus high–energy-density prey (e.g. forage fish and salmon) consumed. Estimated requirements were highest in regions where Steller sea lions consumed higher proportions of low–energy-density prey and experienced the highest rates of population decline

Life history and population dynamics of the finetooth shark (Carcharhinus isodon) in the northeastern Gulf of Mexico

The life history and population dynamics of the finetooth shark (Carcharhinus isodon) in the north-eastern Gulf of Mexico were studied by determining age, growth, size-at-maturity, natural mortality, productivity, and elasticity of vital rates of the population. The von Bertalanffy growth model was estimated as Lt=1559 mm TL (1–e–0.24 (t+2.07)) for females and Lt = 1337 mm TL (1–e–0.41 (t+1.39)) for males. For comparison, the Fabens growth equation was also fitted separately to observed size-at-age data, and the fits to the data were found to be similar. The oldest aged specimens were 8.0 and 8.1 yr, and theoretical longevity estimates were 14.4 and 8.5 yr for females and males, respectively. Median length at maturity was 1187 and 1230 mm TL, equivalent to 3.9 and 4.3 yr for males and females, respectively. Two scenarios, based on the results of the two equations used to describe growth, were considered for population modeling and the results were similar. Annual rates of survivorship estimated through five methods ranged from 0.850/yr to 0.607/yr for scenario 1 and from 0.840/yr to 0.590/yr for scenario 2. Productivities were 0.041/yr for scenario 1 and 0.038/yr for scenario 2 when the population level that produces maximum sustain-able yield is assumed to occur at an instantaneous total mortality rate (Z) equaling 1.5 M, and were 0.071/yr and 0.067/yr, when Z=2 M for scenario 1 and 2, respectively. Mean generation time was 6.96 yr and 6.34 yr for scenarios 1 and 2, respectively. Elasticities calculated through simulation of Leslie matrices averaged 12.6% (12.1% for scenario 2) for fertility, 47.7% (46.2% for scenario 2) for juvenile survival, and 39.7% (41.6% for scenario 2) for adult survival. In all, the finetooth shark exhibits life-history and population characteristics intermediate to those of sharks in the small coastal complex and those from some large coastal species, such as the blacktip shark (Carcharhinus limbatus).

Reproduction in the protogynous black grouper Mexico (Mycteroperca bonaci (Poey)) from the southern Gulf of Mexico

An analysis was made of sexual pattern, spawning season, sizes at sexual maturation, and sex change in black grouper (Mycteroperca bonaci) from the southern Gulf of Mexico. Samples were taken between 1996 and 2000, from industrial and small-craft commercial fi sheries, in offshore and inshore waters of the continental shelf of the Yucatan Peninsula (Campeche Bank), including the shallow waters of National Marine Park Alacranes Reef. For all collected specimens (n=1229), sex and maturation condition were determined by histological analysis of the gonads. The offshore sample consisted of 75.1% females, 24.3% males, and 0.6% transitional-stage fish. All individuals collected from inshore waters were females. Gonadal structure and population structure characteristics for Campeche Bank black grouper were consistent with the characteristics of monandric protogynous hermaphrodism for a serranid fish. Sexually active males and females were observed year-round, although ripening females, with stage-III, -IV, and -V vitellogenic oocytes in the ovaries, dominated in samples taken between December and March. In addition, peak occurrence of ripe-running females with hyaline oocytes or postovulatory follicles (or both) in the ovaries was recorded in January and February. A few precocious females began spawning in October and November, and others were still in spawning condition in May and June. Fifty percent maturity of females was attained at 72.1 cm fork length (FL). Median size at sexual inversion was 103.3 cm FL, and 50% of the females measuring 111.4 cm FL had transformed into males. The southern Gulf of Mexico grouper fishery was considered deteriorated and lacked a well-defined management strategy. Results of the present study provide helpful information on black grouper reproduction in this area and could help Mexican authorities choose appropriate management strategies for this fishery, such as minimum size limit, closed fishing season, and protection of spawning aggregations.

Applications in adaptive cluster sampling of Gulf of Alaska rockfish

Adaptive cluster sampling (ACS) has been the subject of many publications about sampling aggregated populations. Choosing the criterion value that invokes ACS remains problematic. We address this problem using data from a June 1999 ACS survey for rockfish, specifically for Pacific ocean perch (Sebastes alutus), and for shortraker (S. borealis) and rougheye (S. aleutianus) rockfish combined. Our hypotheses were that ACS would outperform simple random sampling (SRS) for S. alutus and would be more applicable for S. alutus than for S. borealis and S. aleutianus combined because populations of S. alutus are thought to be more aggregated. Three alternatives for choosing a criterion value were investigated. We chose the strategy that yielded the lowest criterion value and simulated the higher criterion values with the data after the survey. Systematic random sampling was conducted across the whole area to determine the lowest criterion value, and then a new systematic random sample was taken with adaptive sampling around each tow that exceeded the fixed criterion value. ACS yielded gains in precision (SE) over SRS. Bootstrapping showed that the distribution of an ACS estimator is approximately normal, whereas the SRS sampling distribution is skewed and bimodal. Simulation showed that a higher criterion value results in substantially less adaptive sampling with little tradeoff in precision. When time-efficiency was examined, ACS quickly added more samples, but sampling edge units caused this efficiency to be lessened, and the gain in efficiency did not measurably affect our conclusions. ACS for S. alutus should be incorporated with a fixed criterion value equal to the top quartile of previously collected survey data. The second hypothesis was confirmed because ACS did not prove to be more effective for S. borealis-S. aleutianus. Overall, our ACS results were not as optimistic as those previously published in the literature, and indicate the need for further study of this sampling method.