Varpe Øystein

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  • Article
    Timing of Calanus finmarchicus diapause in stochastic environments
    (Elsevier, 2021-09-20) Bandara, Kanchana ; Varpe, Øystein ; Maps, Frédéric ; Ji, Rubao ; Eiane, Ketil ; Tverberg, Vigdis
    In environments with strong seasonality, many herbivorous zooplankton remain active only during the productive season and undergo a period of inactivity and suppressed development termed ‘diapause’ during the unproductive season. The ability to time the diapause entry and exit in response to the seasonality of the environment is thus essential for their survival. However, timing of diapause may become challenging when environmental conditions vary stochastically across shorter and longer timescales, and particularly when zooplankton lack external cues to predict these variations. In this study, we used a novel individual-based model to study the emerging patterns of diapause timing of the high-latitude marine herbivorous copepod Calanus finmarchicus under shorter- (6-h) and longer-term (interannual) environmental stochasticity. The model simulated growth, development, survival and reproduction (income breeding) of a C. finmarchicus population over multiple calendar years and traced the emergence of behavioral responses and life history strategies. The emergent timing of diapause entry and exit were robust to shorter-term environmental stochasticity, which was manifested through morphological (i.e., body and energy reserve sizes) and behavioral plasticity (i.e., diel vertical migration). Longer-term stochastic variations of temperature and food environments altered the timing of diapause entry, which occurred earlier in warmer years with higher growth potential and vice versa. Irrespective of the modelled environmental variability, diapause exit occurred asynchronously throughout the year. This appeared to be a consequence of a diversified bet hedging strategy, where parents spread the starvation mortality risk of ascending to the upper pelagial at food-deprived times of the year among their offspring. This was a potent strategy, particularly in simulations where the timing of the algal bloom varied stochastically between years, since a fraction of the population was present in the upper pelagial year-round and those that coincided with the emergence of the pelagic primary production survived and produced the next generation.
  • Article
    Ecosystem model intercomparison of under-ice and total primary production in the Arctic Ocean
    (John Wiley & Sons, 2016-01-27) Jin, Meibing ; Popova, Ekaterina E. ; Zhang, Jinlun ; Ji, Rubao ; Pendleton, Daniel ; Varpe, Øystein ; Yool, Andrew ; Lee, Younjoo J.
    Previous observational studies have found increasing primary production (PP) in response to declining sea ice cover in the Arctic Ocean. In this study, under-ice PP was assessed based on three coupled ice-ocean-ecosystem models participating in the Forum for Arctic Modeling and Observational Synthesis (FAMOS) project. All models showed good agreement with under-ice measurements of surface chlorophyll-a concentration and vertically integrated PP rates during the main under-ice production period, from mid-May to September. Further, modeled 30-year (1980–2009) mean values and spatial patterns of sea ice concentration compared well with remote sensing data. Under-ice PP was higher in the Arctic shelf seas than in the Arctic Basin, but ratios of under-ice PP over total PP were spatially correlated with annual mean sea ice concentration, with higher ratios in higher ice concentration regions. Decreases in sea ice from 1980 to 2009 were correlated significantly with increases in total PP and decreases in the under-ice PP/total PP ratio for most of the Arctic, but nonsignificantly related to under-ice PP, especially in marginal ice zones. Total PP within the Arctic Circle increased at an annual rate of between 3.2 and 8.0 Tg C/yr from 1980 to 2009. This increase in total PP was due mainly to a PP increase in open water, including increases in both open water area and PP rate per unit area, and therefore much stronger than the changes in under-ice PP. All models suggested that, on a pan-Arctic scale, the fraction of under-ice PP declined with declining sea ice cover over the last three decades.
  • Article
    A high-resolution modeling study on diel and seasonal vertical migrations of high-latitude copepods
    (Elsevier, 2017-12-26) Bandara, Kanchana ; Varpe, Øystein ; Ji, Rubao ; Eiane, Ketil
    Despite diel and seasonal vertical migrations (DVM and SVM) of high-latitude zooplankton have been studied since the late-19th century, questions still remain about the influence of environmental seasonality on vertical migration, and the combined influence of DVM and SVM on zooplankton fitness. Toward addressing these, we developed a model for simulating DVM and SVM of high-latitude herbivorous copepods in high spatio-temporal resolution. In the model, a unique timing and amplitude of DVM and SVM and its ontogenetic trajectory were defined as a vertical strategy. Growth, survival and reproductive performances of numerous vertical strategies hardwired to copepods spawned in different times of the year were assessed by a fitness estimate, which was heuristically maximized by a Genetic Algorithm to derive the optimal vertical strategy for a given model environment. The modelled food concentration, temperature and visual predation risk had a significant influence on the observed vertical strategies. Under low visual predation risk, DVM was less pronounced, and SVM and reproduction occurred earlier in the season, where capital breeding played a significant role. Reproduction was delayed by higher visual predation risk, and copepods that spawned later in the season used the higher food concentrations and temperatures to attain higher growth, which was efficiently traded off for survival through DVM. Consequently, the timing of SVM did not change much from that predicted under lower visual predation risk, but the body and reserve sizes of overwintering stages and the importance of capital breeding diminished. Altogether, these findings emphasize the significance of DVM in environments with elevated visual predation risk and shows its contrasting influence on the phenology of reproduction and SVM, and moreover highlights the importance of conducting field and modeling work to study these migratory strategies in concert.