Update to the Forage Action Team February 16 th 2017

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Drivers of forage population trends and consumption patterns: environmental, spatial and temporal patterns in Chesapeake Bay forage population distributions and predator consumption Update to the Forage Action Team February 16 th 2017 Ryan Woodland 1, Edward Houde 1, Andre Buchheister 2, Robert Latour 3, Carlos Lozano 1, Christopher Sweetman 3 1 Univ. Maryland Center for Environmental Science, Chesapeake Biological Laboratory, Solomons, MD 20688 2 Humboldt State University, Arcata, CA 95521 3 Viginia Institute of Marine Science, Gloucester Point, VA 23062

Project goals Objective 1 Identify environmental gradients associated with spatial and temporal patterns in relative abundance of forage taxa in Chesapeake Bay Objective 2 Explain how spatial and temporal gradients in environmental variables control consumption of forage taxa, and quantify the effect of forage abundance on consumer populations

Spatial scales and areas of interest Baywide Regional 1. Mainstem 2. Tributaries: Patuxent River Potomac River Rappahannock River York River James River

Species/taxa of interest Forage

Species/taxa of interest Forage Consumers Multiple size-classes

Species/taxa of interest Forage Consumers Multiple size-classes Multiple surveys Chesapeake Bay Program Benthic Survey/WQ monitoring survey MD DNR/VIMS juvenile striped bass index seine surveys MD DNR/VIMS trawl surveys CHESFIMS/TIES midwater trawl survey ChesMMaP trawl survey

Project methods Forage Project Objective Indices Patch-scale Large-scale Time-series Environment Delta-GLM GAM/GAMM GLM/GLMM DFA Predators Consumption-Environment Evacuation model GLM DFA Diet-Environment Delta-GAMM Diet-density dependence GLM Models 1. GLM: generalized linear model 2. GAM: generalized additive model 3. Delta-GLM: delta-generalized linear model 4. Delta-GAM(M): delta-generalized additive (mixed) model 5. DFA: dynamic factor analysis

Forage-environment analysis GAMs: patch-scale patterns yy ii = ββ 0 + ββ 1 xx 1 + ββ 2 xx 2 + ss 1 (xx 1 ) + ss 1 (xx 1 ) ββ nn xx nn + ss nn (xx nn ) Taxonomic resolution Species (n=8), Genus/Family (n=5), Functional (n=5) Response variable ln(biomass per square meter[afdw]) Explanatory variables Class: region, year Continuous: dissolved oxygen, salinity, temperature, depth (CBP survey) Interactions (depth*do, depth*salinity, temperature*do) GLMs: large-scale patterns yy ii = ββ 0 + ββ 1 xx 1 + ββ 2 xx 2 ββ nn xx nn Response variable ln(biomass per square meter[afdw]) Explanatory variables (all continuous) Teleconnections/climate indices (NOAA) Susquehanna River flow (USGS) Cumulative 5 C water temperature degree (CBL/VIMS pier dataset) Ordinate DoY at which cumulative 5 C DD > 500 (CBP survey) Chlorophyll concentration (CBP WQ survey) Hypoxic volume (UMichigan)

GAMs: patch-scale modeling results Example: trumpet worm (Pectinaria gouldii) Sms.si.edu Linear term Estimate SE t-value p Day of year (d) -0.00003 0.00002-1.55 0.12 Depth (m) 0.0007 0.00008 7.94 <.0001 DO (mg/l) 0.0009 0.0002 5.90 <.0001 Temperature (C) -0.0003 0.0002-1.79 0.07 Salinity 0.0006 0.00006 11.20 <.0001

GAMs: patch-scale modeling results Pectinaria gouldii (trumpet worm) Response surface: temperature & DO 1 2 3 4 3 4 5 6 2 1 5 6

GAMs: patch-scale modeling results Pectinaria gouldii (trumpet worm) Response surface: depth & salinity 1 2 3 4 3 4 5 6 2 1 5 6

GAMs: patch-scale modeling results Forage-environment analysis 1. Strong differences among invertebrate groups, even within genera (e.g., Macoma spp.)

GAMs: patch-scale modeling results Forage-environment analysis 1. Strong differences among invertebrate groups, even within genera (e.g., Macoma spp.) 2. At level of Functional group, significant linear effect of both Temperature and DO n.s. n.s. +

Functional groups (and Macoma): Modeled effect of DO at different Temperatures on biomass Macoma spp. Razor clams Mainstem Chesapeake Bay Amphipods & Isopods Other bivalves Mysids Polychaetes

GLMs: large-scale modeling results, Baywide model run Forage-environment analysis 1. Model performance evaluated using information criterion AICc 2. Variables most consistently occurring in best performing models: salinity, spring Chla, degree day phenology, large-scale climate patterns Average water quality Degree day variables Climate patterns Group Taxon Salinity Temp DO Chla Cumulative Timing Flow Hypoxia AMO NAO Species Leptocheirus plumulosus -0.58** 0.37* Macoma balthica -0.42 Macoma mitchelli 0.21 0.26 0.71*** Neanthes succinea 0.34 0.41* Pectinaria gouldii 0.38* -0.47* Rangia cuneata 0.45* -0.38 Taxonomic Anthuridae 0.49* -0.29-0.29 Glyceridae -0.5* -0.61** Macoma -0.59 Nereididae 0.33 0.42* Functional Amphipods & Isopods 0.6*** -0.47** Polychaetes 0.38 0.35* 0.58** -0.28 Other bivalves P-values provided for information grey text: p = 0.2-0.1, black text, no asterisk: p = 0.1-0.05, *: p = 0.05-0.01, **: p = 0.01-0.001, ***: p < 0.001

Forage-environment preliminary results (invertebrates) Local densities major forage groups influenced by dissolved oxygen availability and temperature (some groups) Groups such as Macoma spp., razor clams and polychaetes show negative relationships with temperature A positive relationship with water column DO was observed in all functional groups Interannual patterns in density of several forage groups timing of water temperature warming spring phytoplankton production Large-scale climate patterns (positive NAO, negative AMO)

Project timeline (updated to include extension) Logistics and planning Objective 1 Data verification/validation Expanding forage indices 2016 2017 Mr Ap Ma Jn Jl Au Se Oc No De Ja Fe Mr Ap Ma Jn Jl Au Objective 2 Project deliverables Forage abundance vs enviro. Predator consumption vs environment Density dependent foraging Variance dampening Presentations (Forage Act. Team/Fisheries GIT) Reports (quarterly) Scientific presentations (Tidewater x 2) Peer-review manuscript draft

Questions?

Abundance Indices Minimum Trawlable Number (Millions)

Abundance Indices Minimum Trawlable Number (Millions)

Population-level Consumption Striped Bass Summer Flounder Weakfish

Population-level Consumption Striped Bass Summer Flounder Weakfish

Predator-environment & -forage analysis Consumption indices have been updated to include most recent years in ChesMAPP survey Analysis of multivariate diet composition, diet-forage and consumption-environment underway assessing potential for spatial analysis within mainstem of Chesapeake Bay Atlantic Croaker Striped Bass Spot Summer Flounder White Perch Weakfish

GAMs: patch-scale modeling results Taxonomic group: Macoma spp. ChesapeakeBay.net Linear term Estimate SE t-value p Day of year (d) -0.0001 0.0005-0.29 0.7682 Depth (m) -0.007 0.002-3.57 0.0004 DO (mg/l) -0.006 0.003-1.76 0.0778 Temperature (C) -0.002 0.004-0.47 0.6351 Salinity -0.006 0.001-4.49 <.0001

GAMs: patch-scale modeling results Functional group: Amphipod/Isopods Flickr.com/Gutlab Linear term Estimate SE t-value p Day of year (d) -0.0004 0.0002-2.17 0.0299 Depth (m) -0.00002 0.0007-0.03 0.9779 DO (mg/l) 0.007 0.001 5.15 <.0001 Temperature (C) 0.002 0.002 1.43 0.1527 Salinity -0.005 0.0005-10.56 <.0001

GAMs: patch-scale modeling results

GAMs: patch-scale modeling results

GAMs: patch-scale modeling results

JanJuly5_DD Ord_Day_at_500_5DD Jan-July cumulative 5 C Deg Day 2600 2500 2400 2300 2200 2100 2000 1900 1800 1950 1955 1960 1965 1970 1975 1980 1985 1990 1995 2000 2005 2010 2015 140 130 120 110 100 90 80 Ordinate DoY at which cumulative 5 C DD > 500

DATA Forage Biomass (g m-2, AFDW) Biomass (g m-2, AFDW) 0.02 0.018 0.016 0.014 0.012 0.01 0.008 0.006 0.004 0.002 0 0.08 0.07 0.06 0.05 0.04 0.03 0.02 0.01 0 Lower Bay: Amphipods & Isopods 1995 Lower Bay: other bivalves 1997 1999 2001 2003 2005 2007 2009 2011 2013 Biomass (g m-2, AFDW) 2015 Biomass (g m-2, AFDW) 1995 1997 1999 2001 2003 2005 2007 2009 2011 2013 2015 0.35 0.3 0.25 0.2 0.15 0.1 0.05 0 0.09 0.08 0.07 0.06 0.05 0.04 0.03 0.02 0.01 0 1995 Lower Bay: polychaetes Lower Bay: Macoma spp. 1997 1999 2001 2003 2005 2007 2009 2011 2013 2015 1995 1997 1999 2001 2003 2005 2007 2009 2011 2013 2015

Observed air temperatures Monthly mean water temperature ( C) 35 30 25 20 15 10 5 0 Solomons monitoring station SLIM2: 2008-2011 Water Temp = 0.996*Air Temp + 1.192 R² = 0.96 0 5 10 15 20 25 30 Monthly mean air temperature ( C) Solomons, MD, monthly water temperature ( C) Solomons, MD, monthly air temperature ( C) 30 Reconstructed air temperatures 25 20 15 10 5 0-5 0 100 200 300 400 500 600 700 800 Month of time series from January 1950 - December 2015 Estimated water temperatures 30 25 20 15 10 5 0-5 0 100 200 300 400 500 600 700 800 Month of time series from January 1950 - December 2015