Changing distribution and abundance of small pelagics may drive changes in predator distributions, affecting predator availability to fisheries and surveys. However, small pelagic fish are difficult to survey directly, so we developed a novel method of assessing small pelagic fish aggregate abundance via predator diet data. We used piscivore diet data collected from multiple bottom trawl surveys within a Vector Autoregressive Spatio-Temporal (VAST) model to assess trends of small pelagics on the Northeast US shelf. The goal was to develop a spatial “forage index” to inform survey and/or fishery availability in the bluefish (Pomatomus saltatrix) stock assessment. Using spring and fall surveys from 1973-2020, 20 small pelagic groups were identified as major bluefish prey using the diet data. Then, predators were grouped by diet similarity to identify 19 piscivore species with the most similar diet to bluefish in the region. Diets from all 20 piscivores were combined for the 20 prey groups at each surveyed location, and the total weight of small pelagic prey per predator stomach at each location was input into a Poisson-link delta model to estimate expected prey mass per predator stomach. Best fit models included spatial and spatio-temporal random effects, with predator mean length, number of predator species, and sea surface temperature as catchability covariates. Spring and fall prey indices were split into inshore and offshore areas to reflect changing prey availability over time in areas available to the recreational fishery and the bottom trawl survey, and also to contribute to regional ecosystem reporting
"... it is perhaps the most ferocious and bloodthirsty fish in the sea, leaving in its wake a trail of dead and mangled mackerel, menhaden, herring, alewives, and other species on which it preys." (Collette, et al., 2002)
"From Raritan Bay to Rockaway Inlet, we have had a phenomenal bluefish year with lots of bunker and other bait, ultimately leading to an abundance of bluefish." Mid-Atlantic Bluefish Fishery Performance Report, 2021
Can localized predator-prey observations scale to coastwide assessment and management?
Changing distribution and abundance of small pelagics may drive changes in predator distributions, affecting predator availability to fisheries and surveys. However, small pelagic fish are difficult to survey directly, so we developed a novel method of assessing small pelagic fish aggregate abundance via predator diet data. We used piscivore diet data collected from multiple bottom trawl surveys within a Vector Autoregressive Spatio-Temporal (VAST) model to assess trends of small pelagics on the Northeast US shelf. The goal was to develop a spatial “forage index” to inform survey and/or fishery availability in the bluefish (Pomatomus saltatrix) stock assessment. Using spring and fall surveys from 1973-2020, 20 small pelagic groups were identified as major bluefish prey using the diet data. Then, predators were grouped by diet similarity to identify 19 piscivore species with the most similar diet to bluefish in the region. Diets from all 20 piscivores were combined for the 20 prey groups at each surveyed location, and the total weight of small pelagic prey per predator stomach at each location was input into a Poisson-link delta model to estimate expected prey mass per predator stomach. Best fit models included spatial and spatio-temporal random effects, with predator mean length, number of predator species, and sea surface temperature as catchability covariates. Spring and fall prey indices were split into inshore and offshore areas to reflect changing prey availability over time in areas available to the recreational fishery and the bottom trawl survey, and also to contribute to regional ecosystem reporting
Northeast Fisheries Science Center Diet Data Online: https://fwdp.shinyapps.io/tm2020/
Using NEFSC bottom trawl survey diet data from 1973-2021, 20 small pelagic groups were identified as major bluefish prey with 10 or more observations (in descending order of observations): Longfin squids (Doryteuthis formerly Loligo sp.), Anchovy family (Engraulidae), bay anchovy (Anchoa mitchilli), Atlantic butterfish, (Peprilus triachanthus), Cephalopoda, (Anchoa hepsetus), red eye round herring (Etrumeus teres), Sandlance (Ammodytes sp.), scup (Stenotomus chrysops), silver hake (Merluccius bilinearis), shortfin squids (Illex sp.), Atlantic herring (Clupea harengus), Herring family (Clupeidae), Bluefish (Pomatomus saltatrix), silver anchovy (Engraulis eurystole), longfin inshore squid (Doryteuthis pealeii), Atlantic mackerel (Scomber scombrus), flatfish (Pleuronectiformes), weakfish (Cynoscion regalis), and Atlantic menhaden (Brevoortia tyrannus).
Prey categories such as fish unidentified, Osteichthyes, and unidentified animal remains were not included in the prey list. Although unidentified fish and Osteichthyes can comprise a significant portion of bluefish stomach contents, we cannot assume that unidentified fish in other predator stomachs represent unidentified fish in bluefish stomachs.
Image credits: Striped and bay anchovy photo--Robert Aguilar, Smithsonian Environmental Research Center; redeye round herring photo--https://diveary.com ; sandlance photo--Virginia Institute of Marine Science; all others NOAA Fisheries.
Bottom trawls are not designed for efficient sampling of small pelagics. While we can estimate aggregate planktivore biomass for different ecoregions on the Northeast US shelf, estimates have high observation error. Black lines = NEFSC bottom trawl, Red lines = NEAMAP inshore bottom trawl. Orange line on GB Fall indicates significant increasing long term trend.
VAST models two linear predictors for an index: 1. encounter rate, and 2. positive catch (amount in stomach)
A full model for the first linear predictor ρ1 for each observation i can include:
ρ1(i)=β1(ci,ti)+ω∗1(si,ci)+ε∗1(si,ci,ti)+η1(vi,ci)+nk∑k=1λ1(k)Q(i,k)
The full model for the second linear predictor ρ2 has the same structure, estimating β2, ω2, ε2, η2, and λ2 using the observations, categories, locations, times, and covariates.
We modeled aggregate small pelagic prey as a single category, and apply a Poisson-link delta model to estimate expected prey mass per predator stomach as in (Ng, et al., 2021).
VAST model code and documentation: https://github.com/James-Thorson-NOAA/VAST
Spatial and spatio-temporal correlation decay with increasing distance estimated as κ in a Matern function with fixed smoothness and geometric anisotropy (directional correlation, optionally estimated by the model).
Initial model selection consistently supported the inclusion of spatial and spatio-temporal random effects and anisotropy across all datasets: fall, spring, and annual.
Due to uneven "sampling" of Atlantic herring by predators, (Ng, et al., 2021) recommended aggregating across predators to improve the diet-based Atlantic herring biomass index.
Between 1985-2021 there were:
Bluefish diet collection stations, fall Northeast Fisheries Science Center surveys
The figure shows bluefish diet collection stations for fall surveys, 1985-2019.
NEAMAP survey stations with diet collections for piscivores (n = 3838) had a higher proportion with our defined bluefish prey (n = 2418, 63.0015633%).
All size classes of 50 fish predators captured in the NEFSC bottom trawl survey were grouped by diet similarity to identify the size classes of piscivore species with the most similar diet to bluefish in the region. Diet similarity analysis was completed using the Schoener similarity index (@schoener_nonsynchronous_1970; B. Smith, pers. comm.), and is available available via this link on the NEFSC food habits shiny app. The working group evaluated several clustering methods to develop the predator list (see this link with detailed cluster results).
Predators with highest diet similarity to Bluefish from the NEFSC diet database (1973-2020) include Atlantic cod, Atlantic halibut, buckler dory, cusk, fourspot flounder, goosefish, longfin squid, shortfin squid, pollock, red hake, sea raven, silver hake, spiny dogfish, spotted hake, striped bass, summer flounder, thorny skate, weakfish, and white hake. The NEAMAP survey operates closer to shore than the current NEFSC survey. The NEAMAP dataset includes predators sampled by the NEFSC survey and adds two species, Spanish mackerel and spotted sea trout, not captured by the NEFSC survey offshore but included based on working group expert judgement of prey similarity to bluefish. Predator size classes included are listed in Table 2 of the forage fish index working paper at this link.
Image credits: Weakfish and Spanish mackerel-- https://marinefishesofgeorgia.org ; spotted seatrout-- https://fishinginmiami.com ; Sea Raven photo 2/11/2019 11:07:56 AM, Photographer: Andrew J. Martinez, Location: Massachusetts, Stellwagen Bank NMS; all others NOAA Fisheries.
Number of predator species → likely to affect encounter rate
Mean size of predators → likely to affect amount of prey (Ng, et al., 2021)
Sea surface temperature (SST) → likely to affect predator activity and feeding rate encounter rate and amount of prey
Model selection consistently included number of predator species, mean predator size, and SST as covariates using fall, spring, and annual datasets
All piscivore diet collection stations, fall Northeast Fisheries Science Center surveys →
Diets from all 22 piscivores (including bluefish) were combined for the 20 forage fish (bluefish prey) groups at each surveyed location, and the mean weight of forage fish per predator stomach at each location was calculated. Data for each station included station ID, year, season, date, latitude, longitude, vessel, mean bluefish prey weight (g), mean piscivore length (cm), number of piscivore species, and sea surface temperature (degrees C). Because approximately 10% of survey stations were missing in-situ sea water temperature measurements, National Oceanic and Atmospheric Administration Optimum Interpolation Sea Surface Temperature (NOAA OI SST) V2 High Resolution Dataset [@reynolds_daily_2007] data provided by the NOAA PSL, Boulder, Colorado, USA, from their website at https://psl.noaa.gov were used to fill gaps. For survey stations with in-situ temperature measurements, the in-situ measurement was retained. For survey stations with missing temperature data, OI SST was substituted for input into VAST models.
Models were developed combining all data for the year ("Annual") and with separate data for "Spring" (collection months January - June) and "Fall" (collection months July-December) to align with assumptions used in the bluefish stock assessment. Modeled years included 1985-2021 to align with other data inputs in the bluefish stock assessment.
SST is also likely to affect prey distribution, but differently for each prey species. Therefore, SST is not modeled as a density covariate for aggregate small pelagics.
Maps of key areas for Bluefish assessment indices. The full VAST model grid is shown in brown.
Indices for aggregate small pelagics from piscivore stomachs can be calculated for any subset of the full model domain. Bias correction of the resulting indices is then applied (Thorson, et al., 2016).
NEFSC survey strata definitions are built into the VAST northwest-atlantic
extrapolation grid already. We defined additional new strata to address the recreational inshore-offshore 3 mile boundary. The area within and outside 3 miles of shore was defined using the sf
R package as a 3 nautical mile (approximated as 5.556 km) buffer from a high resolution coastline from thernaturalearth
R package. This buffer was then intersected with the current FishStatsUtils::northwest_atlantic_grid
built into VAST and saved using code here. Then, the new State and Federal waters strata were used to split NEFSC survey strata where applicable, and the new full set of strata were used along with a modified function from FishStatsUtils::Prepare_NWA_Extrapolation_Data_Fn
to build a custom extrapolation grid for VAST as described in detail here.
Time series of VAST estimated fall forage indices for input into the bluefish assessment, 1985-2021
VAST estimated Fall forage biomass density →
Time series of VAST estimated fall forage indices for the 2023 State of the Ecosystem report
aggregate forage important in this system with many forage species: Jason Link horrendogram
potential trends in aggregate forage to be compared with trends in zooplankton and environmental drivers
Complex food web, generalist predators
The original "horrendogram" (Link, 2002)
A new bluefish stock assessment was implemented using the Woods Hole Assessment Model (WHAM) (Stock, et al., 2021).
Forage fish indices were explored as covariates on catchability for the fishery independent bottom trawl surveys, but either did not improve the assessment, or the exploratory models did not converge.
However, the application of the forage fish index to the recreational catch per angler catchability was successful when implemented as an autoregressive process over the time-series with WHAM estimating the standard error. The inclusion of the forage fish index improved the fit of all models.
The use of the forage fish index as a covariate on catchability led to an overall decreasing trend in catchability over time. The recreational index is important in scaling the biomass results, and the lower availability at the end of the time-series led to higher biomass estimates from the assessment including forage fish.
WHAM is a state space stock assessment model framework: https://timjmiller.github.io/wham/
The Bigelow index fit with the fall forage fish index did not improve the model fit (AIC), was slightly worse fit and gave identical results The Albatross index fit with the fall forage fish index did not converge or hessian was not positive definite for any of the models (even when how = 0 for some of them). The MRIP index fit with the annual forage fish index did not converge or hessian was not positive definite for any of the models
Collette, B. B. et al. (2002). Bigelow and Schroeder's Fishes of the Gulf of Maine, Third Edition. 3rd ed. edition. Washington, DC: Smithsonian Books. ISBN: 978-1-56098-951-6.
Deroba, J. J. et al. (2018). "The dream and the reality: meeting decision-making time frames while incorporating ecosystem and economic models into management strategy evaluation". In: Canadian Journal of Fisheries and Aquatic Sciences. ISSN: 0706-652X. DOI: 10.1139/cjfas-2018-0128. URL: http://www.nrcresearchpress.com/doi/10.1139/cjfas-2018-0128 (visited on Jul. 20, 2018).
Link, J. (2002). "Does food web theory work for marine ecosystems?" En. In: Marine Ecology Progress Series 230, pp. 1-9. ISSN: 0171-8630, 1616-1599. DOI: 10.3354/meps230001. URL: https://www.int-res.com/abstracts/meps/v230/p1-9/ (visited on Nov. 04, 2022).
Ng, E. L. et al. (2021). "Predator stomach contents can provide accurate indices of prey biomass". In: ICES Journal of Marine Science 78.3, pp. 1146-1159. ISSN: 1054-3139. DOI: 10.1093/icesjms/fsab026. URL: https://doi.org/10.1093/icesjms/fsab026 (visited on Sep. 01, 2021).
Reynolds, R. W. et al. (2007). "Daily High-Resolution-Blended Analyses for Sea Surface Temperature". EN. In: Journal of Climate 20.22. Publisher: American Meteorological Society Section: Journal of Climate, pp. 5473-5496. ISSN: 0894-8755, 1520-0442. DOI: 10.1175/2007JCLI1824.1. URL: https://journals.ametsoc.org/view/journals/clim/20/22/2007jcli1824.1.xml (visited on Aug. 01, 2022).
Stock, B. C. et al. (2021). "The Woods Hole Assessment Model (WHAM): A general state-space assessment framework that incorporates time- and age-varying processes via random effects and links to environmental covariates". En. In: Fisheries Research 240, p. 105967. ISSN: 0165-7836. DOI: 10.1016/j.fishres.2021.105967. URL: https://www.sciencedirect.com/science/article/pii/S0165783621000953 (visited on May. 26, 2021).
Thorson, J. T. (2019). "Guidance for decisions using the Vector Autoregressive Spatio-Temporal (VAST) package in stock, ecosystem, habitat and climate assessments". En. In: Fisheries Research 210, pp. 143-161. ISSN: 0165-7836. DOI: 10.1016/j.fishres.2018.10.013. URL: http://www.sciencedirect.com/science/article/pii/S0165783618302820 (visited on Feb. 24, 2020).
Thorson, J. T. et al. (2017). "Comparing estimates of abundance trends and distribution shifts using single- and multispecies models of fishes and biogenic habitat". In: ICES Journal of Marine Science 74.5, pp. 1311-1321. ISSN: 1054-3139. DOI: 10.1093/icesjms/fsw193. URL: https://doi.org/10.1093/icesjms/fsw193 (visited on Nov. 04, 2021).
Thorson, J. T. et al. (2016). "Implementing a generic method for bias correction in statistical models using random effects, with spatial and population dynamics examples". En. In: Fisheries Research 175, pp. 66-74. ISSN: 0165-7836. DOI: 10.1016/j.fishres.2015.11.016. URL: https://www.sciencedirect.com/science/article/pii/S0165783615301399 (visited on Jul. 29, 2022).
Slides available at https://noaa-edab.github.io/presentations
Contact: Sarah.Gaichas@noaa.gov
"... it is perhaps the most ferocious and bloodthirsty fish in the sea, leaving in its wake a trail of dead and mangled mackerel, menhaden, herring, alewives, and other species on which it preys." (Collette, et al., 2002)
"From Raritan Bay to Rockaway Inlet, we have had a phenomenal bluefish year with lots of bunker and other bait, ultimately leading to an abundance of bluefish." Mid-Atlantic Bluefish Fishery Performance Report, 2021
Can localized predator-prey observations scale to coastwide assessment and management?
Changing distribution and abundance of small pelagics may drive changes in predator distributions, affecting predator availability to fisheries and surveys. However, small pelagic fish are difficult to survey directly, so we developed a novel method of assessing small pelagic fish aggregate abundance via predator diet data. We used piscivore diet data collected from multiple bottom trawl surveys within a Vector Autoregressive Spatio-Temporal (VAST) model to assess trends of small pelagics on the Northeast US shelf. The goal was to develop a spatial “forage index” to inform survey and/or fishery availability in the bluefish (Pomatomus saltatrix) stock assessment. Using spring and fall surveys from 1973-2020, 20 small pelagic groups were identified as major bluefish prey using the diet data. Then, predators were grouped by diet similarity to identify 19 piscivore species with the most similar diet to bluefish in the region. Diets from all 20 piscivores were combined for the 20 prey groups at each surveyed location, and the total weight of small pelagic prey per predator stomach at each location was input into a Poisson-link delta model to estimate expected prey mass per predator stomach. Best fit models included spatial and spatio-temporal random effects, with predator mean length, number of predator species, and sea surface temperature as catchability covariates. Spring and fall prey indices were split into inshore and offshore areas to reflect changing prey availability over time in areas available to the recreational fishery and the bottom trawl survey, and also to contribute to regional ecosystem reporting
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