1. An adjoint data assimilation approach Physical and Biological Controls on Calanus finmarchicus in the Georges Bank Region GlOBEC broad-scale surveysAcadia (Lynch et al., 1996, 1998)

2. Climatological C. f. distributions (GLOBEC, 1995-1999) Jan: abundances are low (C3<C2)<C5 Feb: reproduction starts; C2>C3>C4>c5; centers are advected along the bank Mar-Apr: abundances are high May: C2,C3 abundances decline Jun: abundances drop sharply on the crest; high centers retain near the Southern Flank (C2, C3) and along the periphery of the bank (C4,C5); high centers retain near the Southern Flank (C2, C3) and along the periphery of the bank (C4,C5); C5>C4>>C3>C2 C5>C4>>C3>C2

3. Questions Where are the off-bank sources in late winter? Where are the off-bank sources in late winter? How are the off-bank sources imported to the bank to initiate the growing season? How are the off-bank sources imported to the bank to initiate the growing season? How do the biological processes & physical transports result in the observed C. f. distributions? How do the biological processes & physical transports result in the observed C. f. distributions? How do these animals disappear from the crest of the bank? How do these animals disappear from the crest of the bank?

4. R2 F2 F3F4 R5 R3 R4 Molting from C1 and C2 mortality Mortality (R<0) C5 molting, mortality, diapause emergence/ entry Molting Flux Campbell et al., 2001

5. Forward Model: Infer R_i and C_i (t=0) by minimizing: First guess: R_i=0, C_i(t=0)=(Cobs on the bank; 0 off-bank)

6. Observations Model results

7. Inferred initial conditions

8. AbundanceInferred Source/sinksMolting fluxadvection

9. Inferred biological terms Important Source regions: the GB, the Georges Basin, Wilkinson Basin, the Browns banks (Feb-Apr.) Mortality: Mar- April, Southern Flank (Food limitation, Campbell et. al., 2001); May-Jun crest (predation, Bollens et al., 1999 )

10. Inferred C3 controls Net bio-gain/loss Jan-Feb Feb-Mar Mar-Apr Apr-May May-June Advection: loss: Wilkinson, Georges Basins; Gains: northwest crest, Georges Basin and the south tip of GSC. Bio-gain: Feb- Apri ; Bio-loss: June; Feb-Apri: Bio>Adv June decline: Bio-losses;

11. Inferred C5 controls Jan-Feb: Advective convergence (Northern Flank,George s Basin) Feb-Apr: Bio, Adv both are important Bio>Adv June: Bio~Adv Net bio-gain/loss Jan-Feb Feb-Mar Mar-Apr Apr-May May-June

12. Conclusions 1. The Scotian Shelf, GOM are important sources of C4 and C5 in late winter. The convergence of advective C5 flux near the northern periphery of the bank seems to be important to seed the bank. 2. Both biological reactions and physical advection are important for the observed distributions. Physical transports increase from winter to spring.

13. Conclusions 3. Biological gains are major contributors for high abundances in Spring; Biological losses are mainly responsible for the decline of C2-C4 in June, while for C5, both biological loss and advective transport are responsible for the June decline; 4. Molting fluxes largely exceed mortality rates (C2-C3, Jan.-Jun.). 5. Mar.15-Apr.15, mortality is high near the southern Flank; May.15-Jun.15, mortality is high on the crest. 6. Results are inferred from modeled flow fields and data on the GB only. More accurate inference about the controls in the off-bank region needs data in those areas and improved estimate of circulation.

14. Future Work Incorporate more data: N3—C1(pump data)— C2—Adult stages; data in other regions. Incorporate more data: N3—C1(pump data)— C2—Adult stages; data in other regions. Use 3D model to study vertical migration as well as the controls already included. Use 3D model to study vertical migration as well as the controls already included. Explore the connection between interannual, synoptic (e.g., storm driven) physical variability & biological variability. Explore the connection between interannual, synoptic (e.g., storm driven) physical variability & biological variability. Sensitivity study to rank the physical & biological controls Sensitivity study to rank the physical & biological controls Extend the study area to a larger scale (e.g., North Atlantic). Extend the study area to a larger scale (e.g., North Atlantic).

16. Molting Flux (F) C: Concentration; D: stage duration; T: temperature; C F: Food concentration Campbell et al., 2001

17. C2 stage duration T limitedT & Chl limited Chla: OReilly & Zetlin (1996); T (Lynch et al., 1996)

18. Climatological Jan-Feb flow (Lynch et al., 996)

19. Seasonal variations

20. Procedure Choose a best first guess of control vector U. Choose a best first guess of control vector U. Integrate the forward model and calculate the cost function. Integrate the forward model and calculate the cost function. Run the adjoint model to calculate the gradient of J to U. Run the adjoint model to calculate the gradient of J to U. Use a descent algorithm to find a new value of U. Use a descent algorithm to find a new value of U. Repeat from the second step until a satisfactory solution has been achieved. Repeat from the second step until a satisfactory solution has been achieved.

21. The inversion reduces model-data misfit by ~90% unconstrained constrained

22. Computed tendencies Increasing season: Jan- Apr; Decline season: Apr-Jun; Centers are located along the bank periphery with advective signatures.

23. Convergence of C5 advective flux (Jan.15)

24. Inferred C2 controls Jan-Feb: biological gains first appears near the Northern edge of the Bank. Feb-May Biological gains/loses and advective transport are important June: bio.-lose is mainly responsible for the decline

25. Inferred C4 controls Net bio-gain/loss Jan-Feb Feb-Mar Mar-Apr Apr-May May-June High abundances in spring: bio-gains; June- decline: bio-losses.

26. Hypotheses Georges Bank populations limited by C. f. supply from GOM diapause stock during winter. Georges Bank populations limited by C. f. supply from GOM diapause stock during winter. GB dynamics influenced by growth conditions on the bank during spring (food limited in April). GB dynamics influenced by growth conditions on the bank during spring (food limited in April). GOM sources to GB become more and more important during spring and summer. GOM sources to GB become more and more important during spring and summer. Courtesy of http://globec.whoi.edu, phase1 projectshttp://globec.whoi.edu

27. MARMAP Observations(1977-1987) Courtesy of www.at-sea.org

28. Figure 4. Estimates of predation impact by 5 predators, based on feeding rates and abundances of predators and prey from Broad Scale Survey samples. Impact is expressed as percent of the stocks of Calanus and Pseudocalanus removed daily, assuming that the predators are non-selective (E=0), and consume these species in the same proportions as they occur. Prey of Clytia are nauplii, prey of the other predators are copepodites and adults (Bollens et. al., online.sfsu.edu).