Frederick S. Scharf

Inferring ecological relationships from the edges of scatter diagrams: Comparison of regression techniques

Scatter diagrams have historically proved useful in the study of associative relationships in ecology. Several important ecological questions involve correlations be- tween variables resulting in polygonal shapes. Two examples that have received consid- erable attention are patterns between prey size and predator size in animal populations and the relationship between animal abundance and body size. Each is typically illustrated using scatter diagrams with upper and lower boundaries of response variables often changing at different rates with changes in the independent variables. Despite recent statistical contri- butions that have stimulated an interest in characterizing the limits of a variable, a consensus on an appropriate methodology to quantify the boundaries of scatter diagrams has not yet been achieved. We tested regression techniques based on least squares and least absolute values models using several independent data sets on prey length and predator length for piscivorous fishes and compared estimated slopes for consistency. Our results indicated that least squares regression techniques were particularly sensitive to outlying y values and irregularities in the distribution of observations, and that they frequently produced incon- sistent estimates of slope for upper and lower bounds. In contrast, quantile regression techniques based on least absolute values models appeared robust to outlying y values and sparseness within data sets, while providing consistent estimates of upper and lower bound slopes. Moreover, the use of quantile regression eliminated the need for an excess of arbitrary decision-making on the part of the investigator. We recommend quantile regression as an improvement to currently available techniques used to examine potential ecological relationships dependent upon quantitative information on the boundaries of polygonal re- lationships.

Vulnerability of marine forage fishes to piscivory: effects of prey behavior on susceptibility to attack and capture

We conducted a series of size-structured laboratory experiments to quantify and compare the susceptibility of several estuarine and marine forage fishes to attack and capture by piscivorous predators. Size-dependent estimates of capture success, handling time, and prey profitability were generated from single-species experiments offering bay anchovy, Atlantic menhaden, Atlantic silverside, and age-0 striped bass to piscivores. Bay anchovy and Atlantic menhaden were most susceptible to capture and yielded high profitability compared to Atlantic silverside and age-0 striped bass prey. Variation in capture success among forage species was particularly influential in generating disparate profitability functions. Although morphological differences among forage species contributed to variation in susceptibility to predation, behavioral analyses indicated that variable reaction distances to approaching predators and activity levels of prey may explain a large fraction of the observed differences in susceptibility. When several forage species were offered to predators simultaneously in larger enclosures, mortality was highest and occurred earlier for bay anchovy and Atlantic menhaden compared to other prey, which points to the strong influence of predator capture success on overall forage fish vulnerability. Our results demonstrate species-specific differences among forage fishes in the ability to avoid attack and capture by piscivores, and we conclude that the

Estimating piscine prey size from partial remains: testing for shifts in foraging mode by juvenile bluefish

Knowledge of prey sizes consumed by a predator aids in the estimation of predation impact. Young-of-the-year bluefish, Pomatomus saltatrix, attack their prey tail-first and often bite their prey in half; this poses a unique problem in determining prey sizes from stomach content analysis. We developed a series of linear regressions to estimate original prey lengths from measurements of eye diameter and caudal peduncle depth for striped bass, Morone saxatilis, bay anchovy, Anchoa mitchilli, American shad, Alosa sapidissima, blueback herring, Alosa aestivalis, Atlantic silverside, Menidia menidia, and white perch, Morone americana. We then used these regressions to estimate original prey sizes from pieces of prey found in stomachs of bluefish collected in the Hudson River estuary from 1990–1993. Lengths of prey that were swallowed whole were compared to estimated lengths of prey that were consumed in pieces. Lengths of prey that were consumed in pieces were larger than prey that were consumed whole. We determined the prey length/predator length ratio at which bluefish began shifting from swallowing their prey whole to partial consumption. Shifting occurred at a ratio of approximately 0.35 irrespective of prey species, suggesting that prey length plays an important role in predator foraging decisions and may contribute to gape limitations. Shifts in foraging mode effectively reduce gape limitation and allow bluefish to consume larger prey sizes which may increase their effect on prey populations.

Recruitment of Juvenile Red Drum in North Carolina: Spatiotemporal Patterns of Year‐Class Strength and Validation of a Seine Survey

Abstract Sixteen years of data from a North Carolina Division of Marine Fisheries seining survey were analyzed to assess the validity of the survey, describe spatial and temporal patterns in recruitment of red drum Sciaenops ocellatus, and evaluate factors that potentially contribute to recruitment variation. Overall, 11,817 age-0 red drum (total length = 11–104 mm) were collected in the survey during 1991–2006. Annual estimates of red drum recruitment were variable over time and showed no long-term trend. The dome-shaped catch-per-unit-effort (CPUE) pattern of all annual cohorts indicated that the seine survey captured peak abundance levels each year; the spatial coverage of the survey also appeared to be adequate. Regional correlations within North Carolina were strongest for areas that were adjacent to one another, and the North Carolina statewide index was not correlated with indices from Florida or Texas, suggesting that factors controlling recruitment of red drum operate at a scale of tens to hundre...

Estuarine Recruitment, Growth, and First-Year Survival of Juvenile Red Drum in North Carolina

Abstract For many marine and estuarine fishes, there is growing evidence that processes occurring during the juvenile life stage can contribute to determining the relative success of recruitment of annual cohorts into the fishery. We studied two consecutive annual cohorts of juvenile red drum Sciaenops ocellatus from estuarine arrival through age 1 in two estuarine systems of southeastern North Carolina. For age-0 cohorts, relative abundance generally peaked in mid to late fall, and fish were present in shallow-water habitats through December before presumably moving to deeper waters during winter. Reappearance of red drum in shallow-water habitats during spring occurred between late March and late April depending on spring warming rates. Considerable differences in relative abundance of age-0 red drum were observed between estuaries; such differences may be related to variations in estuarine hydrology and flushing rates, which probably affect larval delivery and distribution. Otolith-derived hatch date d...

Fishing Mortality in North Carolina's Southern Flounder Fishery: Direct Estimates of Instantaneous Fishing Mortality from a Tag Return Experiment

Abstract Estimation of harvest rates is often a critical component of fishery stock assessment and management. These assessments are often based on catch-at-age data sets generated over many years, but estimates of instantaneous fishing mortality (F) can also be obtained from a shorter-term tag return study. We conducted a 2-year tag return experiment to generate direct estimates of F for southern flounder Paralichthys lethostigma in a North Carolina estuary. The southern flounder supports lucrative commercial and recreational fisheries within the state and has experienced heavy fishing pressure for more than a decade. During 2005 and 2006, fish were captured and tagged with the assistance of commercial harvesters in the New River estuary. Tag returns were used to generate monthly estimates of F, which demonstrated a clear seasonal pattern that was consistent between years. Several important assumptions of the tag return model were accounted for through the use of double-tagged individuals, the distribution of both high- and standard-reward tags, and the completion of an independent controlled experiment to evaluate mortality related to tagging. Annual estimates of F exceeded the short-term management target in both years. Residual patterns suggest that the estimates may actually have been biased low, possibly due to delayed mixing of tagged fish. Thus, despite recently amended fishery regulations, F in the North Carolina southern flounder gill-net fishery still has the potential to greatly exceed targeted levels, which may delay stock recovery. Tag return studies can provide reliable (and nearly real-time) information about F and natural mortality as long as the experimental design addresses specific assumptions related to tagging-induced mortality, tag shedding, and nonreporting of tags.

Effects of Variable Prey and Cohort Dynamics on Growth of Young-of-the-Year Estuarine Bluefish: Evidence for Interactions between Spring- and Summer-Spawned Cohorts

Abstract Previous field studies of bluefish Pomatomus saltatrix have documented variation in young-of-the-year (age-0) growth rates among years and between spring- and summer-spawned cohorts. However, the potential factors responsible for generating variable growth in age-0 bluefish have not been investigated. We constructed an individual-based model that combined size-dependent bluefish foraging with a bioenergetics model to quantify the potential effects of variable prey fish dynamics on first-summer growth of juvenile bluefish. We used long-term monitoring data to define baseline conditions and calibrate the model. We then performed three simulation experiments designed to assess the effects of initial density and arrival timing of prey species and bluefish cohorts on bluefish length distributions on October 1. Simulation experiments indicated that spring-spawned bluefish were robust to fluctuations in prey dynamics because of a spawning strategy that ensures temporal overlap with a diversity of prey f...

Telemetry-Based Mortality Estimates of Juvenile Spot in Two North Carolina Estuarine Creeks

Abstract We estimated natural mortality rates (M) of age-1 Spot Leiostomus xanthurus by using a sonic telemetry approach. Sonic transmitters were surgically implanted into a total of 123 age-1 Spot in two North Carolina estuarine creeks during spring 2009 and 2010, and the fish were monitored by using a stationary acoustic receiver array and manual tracking. Fates of telemetered Spot were inferred based on telemetry information from estimated locations and swimming speeds. Potential competitors of age-1 Spot were assessed through simultaneous otter trawl sampling, while potential predators of Spot were collected using gill nets and trammel nets. The number of inferred natural mortalities was zero in 2009 (based on 29 telemetered Spot at risk) and four in 2010 (based on 52 fish at risk), with fish being at risk for up to about 70 d each year. Catches of potential competitors or predators did not differ between years, and age-1 Spot were not found in analyzed stomach contents of potential predators. Our est...

Histological Analysis Reveals Larger Size at Maturity for Southern Flounder with Implications for Biological Reference Points

Abstract For fish stocks managed using biological reference points based upon spawning biomass, it is critical to have accurate maturity schedules. We investigated size- and age-dependent patterns in maturity for southern flounder Paralichthys lethostigma, a flatfish supporting valuable coastal fisheries in North Carolina and throughout its range. We evaluated both macroscopic and histological methods over two consecutive reproductive seasons. Histological analyses revealed that maturity occurred at larger sizes and older ages than previously estimated. Length at 50% maturity (L 50) was estimated at 408 mm total length (TL), which was more than 60 mm larger than currently assumed, and was relatively stable between study years. We found that only 44% of age-1 southern flounder were mature compared with an estimated 74% in an earlier study. We suspect that most of the differences in maturity timing of southern flounder between our findings and previous studies stem from macroscopic assignment error. During this study, only 61% of fish staged macroscopically as developing were found to be mature based on histological analysis. Assuming incorrectly that all of these fish were mature would have resulted in an L 50 of 375 mm TL, which is closer to previous estimates. Analysis of spawning stock biomass per recruit demonstrated that biological reference points (e.g., F SPR) could be affected considerably by shifting maturity schedules, and the effects could be magnified at larger sizes at entry and higher harvest rates. Given the life history strategy of southern flounder and the lack of a developed offshore fishery or sampling program, which combine to prevent access to fish on the spawning grounds, it is probably most judicious to routinely analyze reproductive tissue samples histologically to ensure accurate information on the timing of maturity.

Factors Affecting Catch-and-Release Mortality of Bluefish

Abstract Recreational harvests of bluefish Pomatomus saltatrix along the U.S. coast from Maine to Florida exceed commercial harvests, and in recent years about 60–70% of angler-captured fish are released alive. The proportion of fish that survive hooking, handling, and release back to the ocean is unknown; however, if catch-and-release mortality is high, it may represent a significant component of the overall bluefish mortality rate. We estimated long-term (21-d) hooking mortality rates of field-captured bluefish and investigated the effects of various factors on postrelease mortality. Age, length, and the occurrence of bleeding were significant factors associated with catch-and-release mortality, which we estimated to be 38.8%. About 65% of the mortality was initial mortality, and the remainder was delayed mortality. We also performed a laboratory study to examine the physiological response of bluefish to two independent processes (hooking and release versus transfer in coolers to the laboratory) relevan...