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.

The state of coral reef ecosystems of the United States and Pacific Freely Associated States: 2008

In the past decade, increased awareness regarding the declining condition of U.S. coral reefs has prompted various actions by governmental and non-governmental organizations. Presidential Executive Order 13089 created the U.S. Coral Reef Task Force (USCRTF) in 1998 to coordinate federal and state/territorial activities (Clinton, 1998), and the Coral Reef Conservation Act of 2000 provided Congressional funding for activities to conserve these important ecosystems, including mapping, monitoring and assessment projects carried out through the support of NOAA’s CRCP. Numerous collaborations forged among federal agencies and state, local, non-governmental, academic and private partners now support a variety of monitoring activities. This report shares the results of many of these monitoring activities, relying heavily on quantitative, spatially-explicit data that has been collected in the recent past and comparisons with historical data where possible. The success of this effort can be attributed to the dedication of over 270 report contributors who comprised the expert writing teams in the jurisdictions and contributed to the National Level Activities and National Summary chapters. The scope and content of this report are the result of their dedication to this considerable collaborative effort. Ultimately, the goal of this report is to answer the difficult but vital question: what is the condition of U.S. coral reef ecosystems? The report attempts to base a response on the best available science emerging from coral reef ecosystem monitoring programs in 15 jurisdictions across the country. However, few monitoring programs have been in place for longer than a decade, and many have been initiated only within the past two to five years. A few jurisdictions are just beginning to implement monitoring programs and face challenges stemming from a lack of basic habitat maps and other ecosystem data in addition to adequate training, capacity building, and technical support. There is also a general paucity of historical data describing the condition of ecosystem resources before major human impacts occurred, which limits any attempt to present the current conditions within an historical context and contributes to the phenomenon of shifting baselines (Jackson, 1997; Jackson et al., 2001; Pandolfi et al., 2005).

Examination of the foraging habits of Pacific harbor seal (Phoca vitulina richardsi) to describe their use of the Umpqua River, Oregon, and their predation on salmonids

The increase in harbor seal (Phoca vitulina richardsi) abundance, concurrent with the decrease in salmonid (Oncorhynchus spp.) and other fish stocks, raises concerns about the potential negative impact of seals on fish populations. Although harbor seals are found in rivers and estuaries, their presence is not necessarily indicative of exclusive or predominant feeding in these systems. We examined the diet of harbor seals in the Umpqua River, Oregon, during 1997 and 1998 to indirectly assess whether or not they were feeding in the river. Fish otoliths and other skeletal structures were recovered from 651 scats and used to identify seal prey. The use of all diagnostic prey structures, rather than just otoliths, increased our estimates of the number of taxa, the minimum number of individuals and percent frequency of occurrence (%FO) of prey consumed. The %FO indicated that the most common prey were pleuronectids, Pacific hake (Merluccius productus), Pacific stag-horn sculpin (Leptocottus armatus), osmerids, and shiner surfperch (Cymatogaster aggregata). The majority (76%) of prey were fish that inhabit marine waters exclusively and fish found in marine and estuarine areas (e.g. anadromous spp.) which would indicate that seals forage predominantly at sea and use the estuary for resting and opportunistic feeding. Salmonid remains were encountered in 39 samples (6%); two samples contained identifiable otoliths, which were determined to be from chi-nook salmon (O. tshawytscha). Because of the complex salmonid composition in the Umpqua River, we used molecular genetic techniques on salmonid bones retrieved from scat to discern species that were rare from those that were abundant. Of the 37 scats with salmonid bones but no otoliths, bones were identified genetically as chinook or coho (O. kisutch) salmon, or steelhead trout (O. mykiss) in 90% of the samples.

Distribution and biology of prowfish (Zaprora silenus) in the northeast Pacific

The prowfish (Zaprora silenus) is an infrequent component of bottom trawl catches collected on stock assessment surveys. Based on presence or absence in over 40,000 trawl catches taken throughout Alaskan waters southward to southern California, prowfish are most frequently encountered in the Gulf of Alaska and the Aleutian Islands at the edge of the continental shelf. Based on data from two trawl surveys, relative abundance indicated by catch per swept area reaches a maximum between 100 m and 200 m depth and is much higher in the Aleutian Islands than in the Gulf of Alaska. Females weigh 3.7% more than males of the same length. Weight-length functions are W (g) = 0.0164 L2.92 (males) and W = 0.0170 L2.92 (females). Length at age does not differ between sexes and is described by L = 89.3(1 – e–0.181(t+0.554)), where L is total length in cm and t is age in years. Females reached 50% maturity at a length of 57.0 cm and an age of 5.1 years. Prowfish diet is almost entirely composed of gelatinous zooplankton, primarily scyphozoa and salps.

Molecular methods for the genetic identification of salmonid prey from Pacific harbor seal (Phoca vitulina richardsi) scat

Twenty-six stocks of Pacific salmon and trout (Oncorhynchus spp.), representing evolutionary significant units (ESU), are listed as threatened or endangered under the Endangered Species Act (ESA) and six more stocks are currently being evaluated for listing. The ecological and economic consequences of these listings are large; therefore considerable effort has been made to understand and respond to these declining populations. Until recently, Pacific harbor seals (Phoca vitulina richardsi) on the west coast increased an average of 5% to 7% per year as a result of the Marine Mammal Protection Act of 1972 (Brown and Kohlman2). Pacific salmon are seasonally important prey for harbor seals (Roffe and Mate, 1984; Olesiuk, 1993); therefore quantifying and understanding the interaction between these two protected species is important for Morphobiologically sound management strategies. Because some Pacific salmonid species in a given area may be threatened or endangered, while others are relatively abundant, it is important to distinguish the species of salmonid upon which the harbor seals are preying. This study takes the first step in understanding these interactions by using molecular genetic tools for species-level identification of salmonid skeletal remains recovered from Pacific harbor seal scats.

Annual estimates of the unobserved incidental kill of pantropical spotted dolphin (Stenella attenuata attenuata) calves in the tuna purse-seine fishery of the eastern tropical Pacific

We estimated the total number of pantropical spotted dolphin (Stenella attenuata) mothers killed without their calves (“calf deficit”) in all tuna purse-seine sets from 1973– 90 and 1996–2000 in the eastern tropical Pacific. Estimates were based on a tally of the mothers killed as reported by color pattern and gender, several color-pattern-based frequency tables, and a weaning model. Over the time series, there was a decrease in the calf deficit from approximately 2800 for the western-southern stock and 5000 in the northeastern stock to about 60 missing calves per year. The mean deficit per set decreased from approximately 1.5 missing calves per set in the mid-1970s to 0.01 per set in the late-1990s. Over the time series examined, from 75% to 95% of the lactating females killed were killed without a calf. Under the assumption that these orphaned calves did not survive without their mothers, this calf deficit represents an approximately 14% increase in the reported kill of calves, which is relatively constant across the years examined. Because the calf deficit as we have defined it is based on the kill of mothers, the total number of missing calves that we estimate is potentially an underestimate of the actual number killed. Further research on the mechanism by which separation of mother and calf occurs is required to obtain better estimates of the unobserved kill of dolphin calves in this fishery.

The dusky rockfishes (Teleostei: Socrpaeniformes) of the North Pacific Ocean: resurrection of Sebastes variabilis (Pallas, 1814) and a redescription of Sebastes ciliatus (Tilesius, 1813)

The dusky rockfish (Sebastes ciliatus) of the North Pacific Ocean has been considered a single variable species with light and dark forms distributed in deep and shallow water, respectively. These forms have been subjected to two distinct fisheries separately managed by federal and state agencies: the light deep form is captured in the offshore trawl fishery; the dark shallow form, in the nearshore jig fishery. The forms have been commonly recognized as the light dusky and dark dusky rockfishes. From morphological evidence correlated with color differences in some 400 specimens, we recognize two species corresponding with these color forms. Sebastes ciliatus (Tilesius) is the dark shallow-water species found in depths of 5−160 m in the western Aleutian Islands and eastern Bering Sea to British Columbia. The name Sebastes variabilis (Pallas) is resurrected from the synonymy of S. ciliatus to apply to the deeper water species known from depths of 12−675 m and ranging from Hokkaido, Japan, through the Aleutian Islands and eastern Bering Sea, to Oregon. Sebastes ciliatus is uniformly dark blue to black, gradually lightening on the ventrum, with a jet black peritoneum, a smaller symphyseal knob, and fewer lateral-line pores compared to S. variabilis. Sebastes variabilis is more variable in body color, ranging from light yellow to a more usual tan or greenish brown to a nearly uniform dark dorsum, but it invariably has a distinct red to white ventrum. Synonymies, diagnoses, descriptions, and geographic distributions are provided for each species.

Diet changes of Pacific cod (Gadus macrocephalus) in Alaska between 1980 and 1995

The diet of Pacific cod (Gadus macrocephalus) in the area of Pavlof Bay, Alaska, was studied in the early 1980s by Albers and Anderson (1985). They found that the dominant prey species were forage species like pandalid shrimp, capelin (Mallotus villosus), and walleye pollock (Theragra chalcogramma). The shrimp fishery in Pavlof Bay began in 1968 and closed in 1980 because of low shrimp abundance (Ruccio and Worton1). Survey data indicate that, during the period between 1972 and 1997, the abundance of forage species such as pandalid shrimp and capelin declined and higher trophic-level groundfish such as Pacific cod increased. There is a general recognition that a long-term ocean climate shift in the Gulf of Alaska has been partially responsible for the observed reorganization of the community structure (Anderson and Piatt, 1999).

Elemental signatures in otoliths of larval walleye pollock (Theragra chalcogramma) from the northeast Pacific Ocean

The population structure of walleye pollock (Theragra chalcogramma) in the northeastern Pacific Ocean remains unknown. We examined elemental signatures in the otoliths of larval and juvenile pollock from locations in the Bering Sea and Gulf of Alaska to determine if there were significant geographic variations in otolith composition that may be used as natural tags of population affinities. Otoliths were assayed by using both electron probe microanalysis (EPMA) and laser ablation inductively coupled plasma mass spectrometry (ICP-MS). Elements measured at the nucleus of otoliths by EPMA and laser ablation ICP-MS differed significantly among locations. However, geographic groupings identified by a multivariate statistical approach from EPMA and ICP-MS were dissimilar, indicating that the elements assayed by each technique were controlled by separate depositional processes within the endolymph. Elemental profiles across the pollock otoliths were generally consistent at distances up to 100 μm from the nucleus. At distances beyond 100 μm, profiles varied significantly but were remarkably consistent among individuals collected at each location. These data may indicate that larvae from various spawning locations are encountering water masses with differing physicochemical properties through their larval lives, and at approximately the same time. Although our results are promising, we require a better understanding of the mechanisms controlling otolith chemistry before it will be possible to reconstruct dispersal pathways of larval pollock based on probe-based analyses of otolith geochemistry. Elemental signatures in otoliths of pollock may allow for the delineation of fine-scale population structure in pollock that has yet to be consistently revealed by using population genetic approaches.

A sudden collapse in distribution of Pacific sardine (Sardinops sagax) off southwestern Australia enables an objective re-assessment of biomass estimates

Fishery-independent estimates of spawning biomass (BSP) of the Pacific sardine (Sardinops sagax) on the south and lower west coasts of Western Australia (WA) were obtained periodically between 1991 and 1999 by using the daily egg production method (DEPM). Ichthyoplankton data collected during these surveys, specifically the presence or absence of S. sagax eggs, were used to investigate trends in the spawning area of S. sagax within each of four regions. The expectation was that trends in BSP and spawning area were positively related. With the DEPM model, estimates of BSP will change proportionally with spawning area if all other variables remain constant. The proportion of positive stations (PPS), i.e., stations with nonzero egg counts — an objective estimator of spawning area — was high for all south coast regions during the early 1990s (a period when the estimated BSP was also high) and then decreased after the mid-1990s. There was a decrease in PPS from the mid-1990s to 1999. The particularly low estimates in 1999 followed a severe epidemic mass mortality of S. sagax throughout their range across southern Australia. Deviations from the expected relationship between BSP and PPS were used to identify uncertainty around estimates of BSP. Because estimation of spawning area is subject to less sampling bias than estimation of BSP, the deviation in the relation between the two provides an objective basis for adjusting some estimates of the latter. Such an approach is particularly useful for fisheries management purposes when sampling problems are suspected to be present. The analysis of PPS undertaken from the same set of samples from which the DEPM estimate is derived will help provide information for stock assessments and for the management of purse-seine fisheries.

Diet of the minimal armhook squid (Berryteuthis anonychus) (Cephalopoda: Gonatidae) in the northeast Pacific during spring

The stomach contents of the minimal armhook squid (Berryteuthis anonychus) were examined for 338 specimens captured in the northeast Pacific during May 1999. The specimens were collected at seven stations between 145−165°W and 39−49°N and ranged in mantle length from 10.3 to 102.2 mm. Their diet comprised seven major prey groups (copepods, chaetognaths, amphipods, euphausiids, ostracods, unidentified fish, and unidentified gelatinous prey) and was dominated by copepods and chaetognaths. Copepod prey comprised four genera, and 86% by number of the copepods were from the genus Neocalanus. Neocalanus cristatus was the most abundant prey taxa, composing 50% by mass and 35% by number of the total diet. Parasagitta elegans (Chaetognatha) occurred in more stomachs (47%) than any other prey taxon. Amphipods occurred in 19% of the stomachs but composed only 5% by number and 3% by mass of the total prey consumed. The four remaining prey groups (euphausiids, ostracods, unidentified fish, and unidentified gelatinous prey) together composed <2% by mass and <1% by number of the diet. There was no major change in the diet through the size range of squid examined and no evidence of cannibalism or predation on other cephalopod species.

The use of parasites in discriminating stocks of Pacific halibut (Hippoglossus stenolepis) in the northeast Pacific

The use of parasites as indicators of the stock structure of Pacific halibut (Hippoglossus stenolepis) in the northeast Pacific was investigated by using 328 adult (>55 cm fork length) halibut from 15 composite localities ranging from northern California to the northern Bering Sea and 96 juvenile (10–55 cm) halibut from five localities ranging from the northern Queen Charlotte Islands to the Bering Sea. Counts of eight selected parasite species (the juvenile acanthocephalans Corynosoma strumosum and C. villosum, the metacestode Nybelinia surmenicola, the digenean metacercaria Otodistomum sp., and the larval nematodes Anisakis simplex, Pseudoterranova decipiens, Contracaecum sp., and Spirurid gen. sp.) that produce infections of long duration, do not multiply in the host, and that have a relatively high abundance in at least one geographic locality were subjected to discriminant function analysis. Juvenile Pacific halibut showed no separation and, even though they were not heavily infected with parasites, the analysis suggested that juveniles could be a mixed stock. Three groups of adults were identified: fish from California to the southern Queen Charlotte Islands, those from the northern Queen Charlotte Islands to the central Bering Sea, and those from the central and north-ern Bering Sea. These groups suggest that the single stock concept be more thoroughly evaluated.

Impact of hatchery releases on the recreational fishery for Pacific threadfin (Polydactylus sexfilis) in Hawaii

The Pacific threadfin (Polydactylus sexfilis) is considered one of the premier Hawaiian food fishes but even with catch limits, seasonal closures, and size limits, catches have declined dramatically since the 1960s. It was identified as the top candidate species for stock enhancement in Hawaii, based on the decline in stocks, high market value, and importance of the fishery. In the stock enhancement program for Pacific threadfin, over 430,000 fingerlings of various sizes were implanted with coded wire tags and released in nursery habitats along the windward coast of Oahu between 1993 and 1998. Because few Pacific threadfin were present in creel surveys conducted between 1994 and 1998, Oahu fishermen were offered a $10 reward for each threadfin that was caught (for both hatchery-reared and wild fish). A total of 1882 Pacific threadfin were recovered from the reward program between March 1998 and May 1999, including 163 hatchery-reared fish, an overall contribution of 8.7% to the fishery. Hatchery-reared fish accounted for as high as 71% of returns in the release areas. Hatchery-reared fish were recovered on average 11.5 km (SD=9.8 km) from the release site, although some had moved as far away as 42 km. Average age for recovered hatchery-reared fish was 495 days; the oldest was 1021 days. Cultured Pacific threadfin juveniles survived and recruited successfully to the recreational fishery, accounting for 10% of fishermen’s catches on the windward side of Oahu. Recruitment to the fishery was highest for the 1997 release year; few juveniles from earlier releases were observed. Presence of a few large, fully developed females in the recreational fishery suggested that hatchery-reared fish can survive, grow, and reproductively contribute to the population. Implementation of an enhancement program that is focused on juveniles and perhaps large females, as part of an integrated fishery management strategy, could speed the recovery of this fish population.