Using social network analysis tools in ecology: Markov process transition models applied to the seasonal trophic network dynamics of the Chesapeake Bay

Ecosystem components interact in complex ways and change over time due to a variety of both internal and external influences (climate change, season cycles, human impacts). Such processes need to be modeled dynamically using appropriate statistical methods for assessing change in network structure. Here we use visualizations and statistical models of network dynamics to understand seasonal changes in the trophic network model described by Baird and Ulanowicz [Baird, D., Ulanowicz, R.E., 1989. Seasonal dynamics of the Chesapeake Bay ecosystem. Ecol. Monogr. 501 (59), 329–364] for the Chesapeake Bay (USA). Visualizations of carbon flow networks were created for each season by using a network graphic analysis tool (NETDRAW). The structural relations of the pelagic and benthic compartments (nodes) in each seasonal network were displayed in a two-dimensional space using spring-embedder analyses with nodes color-coded for habitat associations (benthic or pelagic). The most complex network was summer, when pelagic species such as sea nettles, larval fishes, and carnivorous fishes immigrate into Chesapeake Bay and consume prey largely from the plankton and to some extent the benthos. Winter was the simplest of the seasonal networks, and exhibited the highest ascendency, with fewest nodes present and with most of the flows shifting to the benthic bacteria and sediment POC compartments. This shift in system complexity corresponds with a shift from a pelagic- to benthic-dominated system over the seasonal cycle, suggesting that winter is a mostly closed system, relying on internal cycling rather than external input. Network visualization tools are useful in assessing temporal and spatial changes in food web networks, which can be explored for patterns that can be tested using statistical approaches. A simulation-based continuous-time Markov Chain model called SIENA was used to determine the dynamic structural changes in the trophic network across phases of the annual cycle in a statistical as opposed to a visual assessment. There was a significant decrease in outdegree (prey nodes with reduced link density) and an increase in the number of transitive triples (a triad in which i chooses j and h, and j also chooses h, mostly connected via the non-living detritus nodes in position i), suggesting the Chesapeake Bay is a simpler, but structurally more efficient, ecosystem in the winter than in the summer. As in the visual analysis, this shift in system complexity corresponds with a shift from a pelagic to a more benthic-dominated system from summer to winter. Both the SIENA model and the visualization in NETDRAW support the conclusions of Baird and Ulanowicz [Baird, D., Ulanowicz, R.E., 1989. Seasonal dynamics of the Chesapeake Bay ecosystem. Ecol. Monogr. 501 (59), 329–364] that there was an increase in the Chesapeake Bay ecosystem's ascendancy in the winter. We explain such reduced complexity in winter as a system response to lowered temperature and decreased solar energy input, which causes a decline in the production of new carbon, forcing nodes to go extinct; this causes a change in the structure of the system, making it simpler and more efficient than in summer. It appears that the seasonal dynamics of the trophic structure of Chesapeake Bay can be modeled effectively using the SIENA statistical model for network change.