The biology and ecology of the invasive Indo-Pacific lionfish

The Indo-Pacific lionfishes, Pterois miles and P. volitans, are now established along the Southeast U.S. and Caribbean and are expected to expand into the Gulf of Mexico and Central and South America. Prior to this invasion little was known regarding the biology and ecology of these lionfishes. I provide a synopsis of chronology, taxonomy, local abundance, reproduction, early life history and dispersal, venomology, feeding ecology, parasitology, potential impacts, and possible control and management strategies for the lionfish invasion. This information was collected by review of the literature and by direct field and experimental study. I confirm the existence of an unusual supraocular tentacle phenotype and suggest that the high prevalence of this phenotype in the Atlantic is not the result of selection, but likely ontogenetic change. To describe the trophic impacts of lionfish, I report a comprehensive assessment of diet that describes lionfish as a generalist piscivore that preys on over 40 species of teleost comprising more than 20 families. Next, I use the histology of gonads to describe both oogenesis and reproductive dynamics of lionfish. Lionfish mature relatively early and reproduce several times per month throughout the entire calendar year off North Carolina and the Bahamas. To investigate predation, an important component of natural mortality, I assessed the vulnerability of juvenile lionfish to predation by native serranids. Juvenile lionfish are not readily consumed by serranids, even after extreme periods of starvation. Last, I used a stage-based, matrix population model to estimate the scale of control that would be needed to reduce an invading population of lionfish. Together, this research provides the first comprehensive assessment on lionfish biology and ecology and explains a number of life history and ecological interactions that have facilitated the unprecedented and rapid establishment of this invasive finfish. Future research is needed to understand the scale of impacts that lionfish could cause, especially in coral reef ecosystems, which are already heavily stressed. This research further demonstrates the need for lionfish control strategies and more rigorous prevention and early detection and rapid response programs for marine non-native introductions.

Prevalence of brevetoxins in prey fish of bottlenose dolphins in Sarasota Bay, Florida

Blooms of the brevetoxin-producing dinoflagellate Karenia brevis have been linked to high mortality of bottlenose dolphins Tursiops truncatus on Florida’s Gulf of Mexico coast. A clear understanding of trophic transfer of brevetoxin from its algal source up the food web to top predators is needed to assess exposure of affected dolphin populations. Prey fish constitute a means of accumulating and transferring brevetoxins and are potential vectors of brevetoxin to dolphins frequently exposed to K. brevis blooms. Here we report results of brevetoxin analyses of the primary fish species consumed by long-term resident bottlenose dolphins inhabiting Sarasota Bay, Florida. Fish collected during K. brevis blooms in 2003 to 2006 were analyzed by competitive enzyme-linked immunosorbent assay (ELISA) and had brevetoxin concentrations ranging from 4 to 10844 ng PbTx-3 eq g–1 tissue. Receptor binding assay (RBA) and liquid chromatography–mass spectrometry (LC-MS) analysis confirmed toxicity and the presence of parent brevetoxins and known metabolites. Fish collected in the absence of K. brevis blooms tested positive for brevetoxin by ELISA and RBA, with concentrations up to 1500 ng PbTx-3 eq g–1 tissue. These findings implicate prey fish exposed to K. brevis blooms as brevetoxin vectors for their dolphin predators and provide a critical analysis of persistent brevetoxin loads in the food web of dolphins repeatedly exposed to Florida red tides.

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.

Diet shifts of juvenile red snapper (Lutjanus campechanus) with changes in habitat and fish size

We examined the diets and habitat shift of juvenile red snapper (Lutjanus campechanus) in the northeast Gulf of Mexico. Fish were collected from open sand-mud habitat (little to no relief), and artificial reef habitat (1-m3 concrete or PVC blocks), from June 1993 through December 1994. In 1994, fish settled over open habitat from June to September, as shown by trawl collections, then began shifting to reef habitat — a shift that was almost completed by December as observed by SCUBA visual surveys. Stomachs were examined from 1639 red snapper that ranged in size from 18.0 to 280.0 mm SL. Of these, 850 fish had empty stomachs, and 346 fish from open habitat and 443 fish from reef habitat contained prey. Prey were identified to the lowest possible taxon and quantified by volumetric measurement. Specific volume of particular prey taxa were calculated by dividing prey volume by individual fish weight. Red snapper shifted diets with increasing size. Small red snapper (<60 mm SL) fed mostly on chaetognaths, copepods, shrimp, and squid. Large red snapper (60–280 mm SL) shifted feeding to fish prey, greater amounts of squid and crabs, and continued feeding on shrimp. We compared red snapper diets for overlapping size classes (70–160 mm SL) of fish that were collected from both habitats (Bray-Curtis dissimilarity index and multidimensional scaling analysis). Red snapper diets separated by habitat type rather than fish size for the size ranges that overlapped habitats. These diet shifts were attributed to feeding more on reef prey than on open-water prey. Thus, the shift in habitat shown by juvenile red snapper was reflected in their diet and suggested differential habitat values based not just on predation refuge but food resources as well.

Distribution, age, and growth of young-of-the year greater amberjack (Seriola dumerili) associated with pelagic Sargassum

Patterns of distribution and growth were examined for young-of-the-year (YOY) greater amberjack (Seriola dumerili) associated with pelagic Sargassum in the NW Gulf of Mexico. Seriola dumerili were collected off Galveston, Texas, from May to July over a two-year period (2000 and 2001) in both inshore (<15 nautical miles [nmi]) and offshore zones (15−70 nmi). Relative abundance of YOY S. dumerili (32−210 mm standard length) from purse-seine collections peaked in May and June, and abundance was highest in the offshore zone. Ages of S. dumerili ranged from 39 to 150 days and hatching-date analysis indicated that the majority of spawning events occurred from February to April. Average daily growth rates of YOY S. dumerili for 2000 and 2001 were 1.65 mm/d and 2.00 mm/d, respectively. Intra-annual differences in growth were observed; the late-season (April) cohort experienced the fastest growth in both years. In addition, growth was significantly higher for S. dumerili collected from the offshore zone. Mortality was approximated by using catch-curve analysis, and the predicted instantaneous mortality rate (Z) of YOY S. dumerili was 0.0045 (0.45%/d).

Demographic assessment of the blue crab (Callinectes sapidus) in Chesapeake Bay using extractable lipofuscins as age markers

The blue crab (Callinectes sapidus) plays an important economic and ecological role in estuaries and coastal habitats from the Gulf of Mexico to the east coast of North America, but demographic assessments are limited by length-based methods. We applied an alternative aging method using biochemical measures of metabolic byproducts (lipofuscins) sequestered in the neural tissue of eyestalks to examine population age structure. From Chesapeake Bay, subsamples of animals collected from the 1998–99 (n=769) and 1999–2000 (n=367) winter dredge surveys were collected and lipofuscin was measured. Modal analysis of the lipofuscin index provided separation into three modes, whereas carapace-width data collected among the same individuals showed two broad modes. Lipofuscin modal analysis indicated that most adults (carapace width >120 mm) were <2 years old. The results indicate that use of extractable lipofuscin can provide a more accurate and better resolved estimation of demographic structure of blue crab populations in the field than size alone.

Age and growth estimates of the winter skate (Leucoraja ocellata) in the western Gulf of Maine

Age and growth estimates for the winter skate (Leucoraja ocellata) were estimated from vertebral band counts on 209 fish ranging in size from 145 to 940 mm total length (TL). An index of average percent error (IAPE) of 5.8% suggests that our aging method represents a precise approach to the age assessment of L. ocellata. Marginal increments were significantly different between months (Kruskal-Wallis P<0.001) and a distinct trend of increasing monthly increment growth began in July. Estimates of von Bertalanffy growth parameters suggest that females attain a slightly larger asymptotic TL (L∞=1374 mm) than males (L∞=1218 mm) and grow more slowly (k=0.059 and 0.074, respectively). The oldest ages obtained for the winter skate were 19 years for males and 18 years for females, which corresponded to total lengths of 932 mm and 940 mm, respectively. The results indicate that the winter skate exhibits the characteristics that have made other elasmobranch populations highly susceptible to exploitation by commercial fisheries.

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.

The early life history of swordfish (Xiphias gladius) in the western North Atlantic

Lengths and ages of sword-fish (Xiphias gladius) estimated from increments on otoliths of larvae collected in the Caribbean Sea, Florida Straits, and off the southeastern United States, indicated two growth phases. Larvae complete yolk and oil globule absorption 5 to 6 days after hatching (DAH). Larvae <13 mm preserved standard length (PSL) grow slowly (~0.3 mm/d); larvae from 13 to 115 mm PSL grow rapidly (~6 mm/d). The acceleration in growth rate at 13 days follows an abrupt (within 3 days) change in diet, and in jaw and alimentary canal structure. The diet of swordfish larvae is limited. Larvae <8 mm PSL from the Caribbean, Gulf of Mexico, and off the southeastern United States eat exclusively copepods, primarily of one genus, Corycaeus. Larvae 9 to 11 mm eat copepods and chaetognaths; larvae >11 mm eat exclusively neustonic fish larvae. This diet indicates that young larvae <11 mm occupy the near-surface pelagia, whereas, older and longer larvae are neustonic. Spawning dates for larvae collected in various regions of the western North Atlantic, along with the abundance and spatial distribution of the youngest larvae, indicate that spawning peaks in three seasons and in five regions. Swordfish spawn in the Caribbean Sea, or possibly to the east, in winter, and in the western Gulf of Mexico in spring. Elsewhere swordfish spawn year-round, but spawning peaks in the spring in the north-central Gulf of Mexico, in the summer off southern Florida, and in the spring and early summer off the southeastern United States. The western Gulf Stream frontal zone is the focus of spawning off the southeastern coast of the United States, whereas spawning in the Gulf of Mexico seems to be focused in the vicinity of the Gulf Loop Current. Larvae may use the Gulf of Mexico and the outer continental shelf off the east coast of the United States as nursery areas. Some larvae may be transported northward, but trans-Atlantic transport of larvae is unlikely.

The physiological effects of multiple forced submergences in loggerhead sea turtles (Caretta caretta)

Sea turtles are subjected to involuntary submergence and potential mortality due to incidental capture by the commercial shrimp fishing industry. Despite implementation of turtle excluder devices (TEDs) to reduce at-sea mortality, dead stranded turtles continue to be found in near-record numbers along the coasts of the western Atlantic Ocean and northern Gulf of Mexico. Although this mortality may be due to an increase in the number of turtles available to strand, one alternative explanation is that sea turtles are repetitively submerged (as one fishing vessel follows the path of another) in legal TEDs. In the present study, laboratory and field investigations were undertaken to examine the physiological effects of multiple submergence of loggerhead sea turtles (Caretta caretta). Turtles in the laboratory study were confined during the submersion episodes, whereas under field conditions, turtles were released directly into TED-equipped commercial fishing nets. Under laboratory and field conditions, pre- and postsubmergence blood samples were collected from turtles submerged three times at 7.5 min per episode with an in-water rest interval of 10, 42, or 180 min between submergences. Analyses of pre- and postsubmergence blood samples revealed that the initial submergence produced a severe and pronounced metabolic and respiratory acidosis in all turtles. Successive submergences produced significant changes in blood pH, Pco2, and lactate, although the magnitude of the acid-base imbalance was substantially reduced as the number of submergences increased. In addition, increasing the interval between successive submergences permitted greater recovery of blood homeostasis. No turtles died during these studies. Taken together, these data suggest that repetitive sub-mergence of sea turtles in TEDs would not significantly affect their survival potential provided that the animal has an adequate rest interval at the surface between successive submergences.

Reproduction of blacknose shark (Carcharhinus acronotus) in coastal waters off northeastern Brazil

The blacknose shark, Carcharhinus acronotus, is a relatively small carcharinid, typically inhabiting continental shelf areas in the western Atlantic Ocean, from North Carolina throughout the Gulf of Mexico (Bigelow and Schroeder, 1948) and along the South American coast to Rio de Janeiro (Compagno, 1984). The abundance of this shark in nearshore areas throughout its distribution makes it accessible to commercial fishing, mainly from inshore hook-and-line and gill-net fisheries (Trent et al., 1997; Mattos and Hazin1).

Movement of American lobster (Homarus americanus) in the southwestern Gulf of St. Lawrence

A total of 42,445 American lobsters (Homarus americanus) were tagged in thirty-one sites throughout the southwestern Gulf of St. Lawrence between 1980 and 1997. Results from the recapture of 8503 tagged lobsters showed small distances traveled between the release and the recapture position for animals ranging in size from 51 to 152 mm carapace length. The average distance traveled ranged from 2 km in parts of Baie des Chaleurs and western Cape Breton to 19 km in central Northumberland Strait. Lobsters moved generally along the shore (93% of the dispersion was in areas between the shore and the 20-m bathymetric contour). As a result, lobsters traveled longer distances in sites characterized by a gradually sloping bottom where the distance between the shore and the 20-m contour line was extensive in contrast to areas characterized by rapidly changing depths and by a relatively small amount of habitat shallower than 20 m. In the majority of sites (14 of 19) there was no significant difference between males and females in the average distance they traveled. In four of the five sites females moved farther than males. In general, the average distance traveled by berried females was shorter than that traveled by males or nonberried females. No relationship was observed between the distance traveled and the size of the animal. There was no strong evidence of a relationship between the average distance traveled and the number of days at liberty. In general, lobsters in the southwestern Gulf of St. Lawrence traveled short distances and dispersion was restricted to the nearshore habitat. Further, the distance traveled was not correlated to size, sex, or years at large. These findings show that there is little interaction between American lobsters from different fishing areas at the benthic level and that American lobster movements should have minimal consequences for management of the species in the southwestern Gulf of St. Lawrence.

Shortspine thornyhead (Sebastolobus alascanus) abundance and habitat associations in the Gulf of Alaska

Shortspine thornyhead (Sebastolobus alascanus) abundance was estimated from 107 video transects at 27 stations recorded from a research submersible in 1991 off southeast Alaska at depths ranging from 165 to 355 m. Numbers of invertebrates in seven major taxa were estimated, as was substrate type. Thornyhead abundance ranged from 0 to 7.5/100 m2, with a mean of 1.22/100 m2, and was positively correlated with depth and amount of hard substrate. Invertebrate abundances were not significantly correlated with numbers of thornyheads. Shortspine thornyhead abundance estimates from this study were several times higher than estimates produced by bottom trawl surveys off southeast Alaska in 1990 and 1993, the two years of survey that encompassed the submersible transects; however, the trend of increasing abundance with depth was similar in the trawl surveys and in the submersible transects, suggesting that trawl surveys systematically underestimate abundance of shortspine thornyheads

Physiological ecology of juvenile chinook salmon (Oncorhynchus tshawytscha) at the southern end of their distribution, the San Francisco Estuary and Gulf of the Farallones, California

Juvenile chinook salmon, Oncorhynchus tshawytscha, from natal streams in California’s Central Valley demonstrated little estuarine dependency but grew rapidly once in coastal waters. We collected juvenile chinook salmon at locations spanning the San Francisco Estuary from the western side of the freshwater delta—at the confluence of the Sacramento and San Joaquin Rivers—to the estuary exit at the Golden Gate and in the coastal waters of the Gulf of the Farallones. Juveniles spent about 40 d migrating through the estuary at an estimated rate of 1.6 km/d or faster during their migration season (May and June 1997) toward the ocean. Mean growth in length (0.18 mm/d) and weight (0.02 g/d) was insignificant in young chinook salmon while in the estuary, but estimated daily growth of 0.6 mm/d and 0.5 g/d in the ocean was rapid (P≤0.001). Condition (K factor) declined in the estuary, but improved markedly in ocean fish. Total body protein, total lipid, triacylglycerols (TAG), polar lipids, cholesterol, and nonesterified fatty acids concentrations did not change in juveniles in the estuary, but total lipid and TAG were depleted in ocean juveniles. As young chinook migrated from freshwater to the ocean, their prey changed progressively in importance from invertebrates to fish larvae. Once in coastal waters, juvenile salmon appear to employ a strategy of rapid growth at the expense of energy reserves to increase survival potential. In 1997, environmental conditions did not impede development: freshwater discharge was above average and water temperatures were only slightly elevated, within the species’ tolerance. Data suggest that chinook salmon from California’s Central Valley have evolved a strong ecological propensity for a ocean-type life history. But unlike populations in the Pacific Northwest, they show little estuarine dependency and proceed to the ocean to benefit from the upwelling-driven, biologically productive coastal waters.

Changes over time in the spatial distribution of walleye pollock (Theragra chalcogramma) in the Gulf of Alaska, 1984-1996

Triennial bottom trawl survey data from 1984 to 1996 were used to evaluate changes in the summer distribution of walleye pollock in the western and central Gulf of Alaska. Differences between several age groups of pollock were evaluated. Distribution was examined in relation to several physical characteristics, including bottom depth and distance from land. Interspecies associations were also analyzed with the Bray-Curtis clustering technique to better understand community structure. Our results indicated that although the population numbers decreased, high concentrations of pollock remained in the same areas during 1984–96. However, there was an increase in the number of stations where low-density pollock concentrations of all ages were observed, which resulted in a decrease in mean population density of pollock within the GOA region. Patterns emerging from our data suggested an alternative to Mac-Call’s “basin hypothesis” which states that as population numbers decrease, there should be a contraction of the population range to optimal habitats. During 1984–96 there was a concurrent precipitous decline in Steller sea lions in the Gulf of Alaska. The results of our study suggest that decreases in the mean density of adult pollock, the main food in the Steller sea lion diet, combined with slight changes in the distribution of pollock (age-1 pollock in particular) in the mid-1980s, may have contributed to decreased foraging efficiency in Steller sea lions. Our results support the prevailing conceptual model for pollock ontogeny, although there is evidence that substantial spawning may also occur outside of Shelikof Strait.