Andrew Rassweiler

Analysis of abrupt transitions in ecological systems

The occurrence and causes of abrupt transitions, thresholds, or regime shifts between ecosystem states are of great concern and the likelihood of such transitions is increasing for many ecological systems. General understanding of abrupt transitions has been advanced by theory, but hindered by the lack of a common, accessible, and data-driven approach to characterizing them. We apply such an approach to 30–60 years of data on environmental drivers, biological responses, and associated evidence from pelagic ocean, coastal benthic, polar marine, and semi-arid grassland ecosystems. Our analyses revealed one case in which the response (krill abundance) linearly tracked abrupt changes in the driver (Pacific Decadal Oscillation), but abrupt transitions detected in the three other cases (sea cucumber abundance, penguin abundance, and black grama grass production) exhibited hysteretic relationships with drivers (wave intensity, sea-ice duration, and amounts of monsoonal rainfall, respectively) through a variety of response mechanisms. The use of a common approach across these case studies illustrates that: the utility of leading indicators is often limited and can depend on the abruptness of a transition relative to the lifespan of responsive organisms and observation intervals; information on spatiotemporal context is useful for comparing transitions; and ancillary information from associated experiments and observations aids interpretation of response-driver relationships. The understanding of abrupt transitions offered by this approach provides information that can be used to manage state changes and underscores the utility of long-term observations in multiple sentinel sites across a variety of ecosystems.

Evidence for a general species-time-area relationship

The species-area relationship (SAR) plays a central role in biodiversity re- search, and recent work has increased awareness of its temporal analogue, the species- time relationship (STR). Here we provide evidence for a general species-time-area rela- tionship (STAR), in which species number is a function of the area and time span of sampling, as well as their interaction. For eight assemblages, ranging from lake zooplankton to desert rodents, this model outperformed a sampling-based model and two simpler models in which area and time had independent effects. In every case, the interaction term was negative, meaning that rates of species accumulation in space decreased with the time span of sampling, while species accumulation rates in time decreased with area sampled. Al- though questions remain about its precise functional form, the STAR provides a tool for scaling species richness across time and space, for comparing the relative rates of species turnover in space and time at different scales of sampling, and for rigorous testing of mechanisms proposed to drive community dynamics. Our results show that the SAR and STR are not separate relationships but two dimensions of one unified pattern.

Global regime shift dynamics of catastrophic sea urchin overgrazing

A pronounced, widespread and persistent regime shift among marine ecosystems is observable on temperate rocky reefs as a result of sea urchin overgrazing. Here, we empirically define regime-shift dynamics for this grazing system which transitions between productive macroalgal beds and impoverished urchin barrens. Catastrophic in nature, urchin overgrazing in a well-studied Australian system demonstrates a discontinuous regime shift, which is of particular management concern as recovery of desirable macroalgal beds requires reducing grazers to well below the initial threshold of overgrazing. Generality of this regime-shift dynamic is explored across 13 rocky reef systems (spanning 11 different regions from both hemispheres) by compiling available survey data (totalling 10 901 quadrats surveyed in situ) plus experimental regime-shift responses (observed during a total of 57 in situ manipulations). The emergent and globally coherent pattern shows urchin grazing to cause a discontinuous ‘catastrophic’ regime shift, with hysteresis effect of approximately one order of magnitude in urchin biomass between critical thresholds of overgrazing and recovery. Different life-history traits appear to create asymmetry in the pace of overgrazing versus recovery. Once shifted, strong feedback mechanisms provide resilience for each alternative state thus defining the catastrophic nature of this regime shift. Importantly, human-derived stressors can act to erode resilience of desirable macroalgal beds while strengthening resilience of urchin barrens, thus exacerbating the risk, spatial extent and irreversibility of an unwanted regime shift for marine ecosystems.

Marine protected areas and the value of spatially optimized fishery management

There is a growing focus around the world on marine spatial planning, including spatial fisheries management. Some spatial management approaches are quite blunt, as when marine protected areas (MPAs) are established to restrict fishing in specific locations. Other management tools, such as zoning or spatial user rights, will affect the distribution of fishing effort in a more nuanced manner. Considerable research has focused on the ability of MPAs to increase fishery returns, but the potential for the broader class of spatial management approaches to outperform MPAs has received far less attention. We use bioeconomic models of seven nearshore fisheries in Southern California to explore the value of optimized spatial management in which the distribution of fishing is chosen to maximize profits. We show that fully optimized spatial management can substantially increase fishery profits relative to optimal nonspatial management but that the magnitude of this increase depends on characteristics of the fishing fleet and target species. Strategically placed MPAs can also increase profits substantially compared with nonspatial management, particularly if fishing costs are low, although profit increases available through optimal MPA-based management are roughly half those from fully optimized spatial management. However, if the same total area is protected by randomly placing MPAs, starkly contrasting results emerge: most random MPA designs reduce expected profits. The high value of spatial management estimated here supports continued interest in spatially explicit fisheries regulations but emphasizes that predicted increases in profits can only be achieved if the fishery is well understood and the regulations are strategically designed.

BIOMASS RATHER THAN GROWTH RATE DETERMINES VARIATION IN NET PRIMARY PRODUCTION BY GIANT KELP

Net primary production (NPP) is influenced by disturbance-driven fluctuations in foliar standing crop (FSC) and resource-driven fluctuations in rates of recruitment and growth, yet most studies of NPP have focused primarily on factors influencing growth. We quantified NPP, FSC, recruitment, and growth rate for the giant kelp, Macrocystis pyrifera, at three kelp forests in southern California, U.S.A., over a 54-month period and determined the relative roles of FSC, recruitment, and growth rate in contributing to variation in annual NPP. Net primary production averaged between 0.42 and 2.38 kg dry mass x m(-2) x yr(-1) at the three sites. The initial FSC present at the beginning of the growth year and the recruitment of new plants during the year explained 63% and 21% of the interannual variation observed in NPP, respectively. The previous years NPP and disturbance from waves collectively accounted for 80% of the interannual variation in initial FSC. No correlation was found between annual growth rate (i.e., the amount of new kelp mass produced per unit of existing kelp mass) and annual NPP (i.e., the amount of new kelp mass produced per unit area of ocean bottom), largely because annual growth rate was consistent compared to initial FSC and recruitment, which fluctuated greatly among years and sites. Although growth rate was a poor predictor of variation in annual NPP, it was principally responsible for the high mean values observed for NPP by Macrocystis. These high mean values reflected rapid growth (average of approximately 2% per day) of a relatively small standing crop (maximum annual mean = 444 g dry mass/m2) that replaced itself approximately seven times per year. Disturbance-driven variability in FSC may be generally important in explaining variation in NPP, yet it is rarely examined because cycles of disturbance and recovery occur over timescales of decades or more in many systems. Considerable insight into how variation in FSC drives variation in NPP may be gained by studying systems such as giant kelp forests that are characterized by frequent disturbance and rapid rates of growth and recruitment.

The value of spatial information in MPA network design

The science of spatial fisheries management, which combines ecology, oceanography, and economics, has matured significantly. As a result, there have been recent advances in exploiting spatially explicit data to develop spatially explicit management policies, such as networks of marine protected areas (MPAs). However, when data are sparse, spatially explicit policies become less viable, and we must instead rely on blunt policies such as total allowable catches or imprecisely configured networks of MPAs. Therefore, spatial information has the potential to change management approaches and thus has value. We develop a general framework within which to analyze the value of information for spatial fisheries management and apply that framework to several US Pacific coast fisheries. We find that improved spatial information can increase fishery value significantly (>10% in our simulations), and that it changes dramatically the efficient management approach—switching from diffuse effort everywhere to a strategy where fishing is spatially targeted, with some areas under intensive harvest and others closed to fishing. Using all available information, even when incomplete, is essential to management success and may as much as double fishery value relative to using (admittedly incorrect) assumptions commonly invoked.

Wave disturbance overwhelms top-down and bottom-up control of primary production in California kelp forests.

We took advantage of regional differences in environmental forcing and consumer abundance to examine the relative importance of nutrient availability (bottom-up), grazing pressure (top-down), and storm waves (disturbance) in determining the standing biomass and net primary production (NPP) of the giant kelp Macrocystis pyrifera in central and southern California. Using a nine-year data set collected from 17 sites we show that, despite high densities of sea urchin grazers and prolonged periods of low nutrient availability in southern California, NPP by giant kelp was twice that of central California where nutrient concentrations were consistently high and sea urchins were nearly absent due to predation by sea otters. Waves associated with winter storms were consistently higher in central California, and the loss of kelp biomass to winter wave disturbance was on average twice that of southern California. These observations suggest that the more intense wave disturbance in central California limited NPP by giant kelp under otherwise favorable conditions. Regional patterns of interannual variation in NPP were similar to those of wave disturbance in that year-to-year variation in disturbance and NPP were both greater in southern California. Our findings provide strong evidence that regional differences in wave disturbance overwhelmed those of nutrient supply and grazing intensity to determine NPP by giant kelp. The important role of disturbance in controlling NPP revealed by our study is likely not unique to giant kelp forests, as vegetation dynamics in many systems are dominated by post-disturbance succession with climax communities being relatively uncommon. The effects of disturbance frequency may be easier to detect in giant kelp because it is fast growing and relatively short lived, with cycles of disturbance and recovery occurring on time scales of years. Much longer data sets (decades to centuries) will likely be needed to properly evaluate the role of disturbance relative to other processes in determining patterns of NPP in other systems.

Global patterns of kelp forest change over the past half-century

Significance Kelp forests support diverse and productive ecological communities throughout temperate and arctic regions worldwide, providing numerous ecosystem services to humans. Literature suggests that kelp forests are increasingly threatened by a variety of human impacts, including climate change, overfishing, and direct harvest. We provide the first globally comprehensive analysis of kelp forest change over the past 50 y, identifying a high degree of variation in the magnitude and direction of change across the geographic range of kelps. These results suggest region-specific responses to global change, with local drivers playing an important role in driving patterns of kelp abundance. Increased monitoring aimed at understanding regional kelp forest dynamics is likely to prove most effective for the adaptive management of these important ecosystems. Kelp forests (Order Laminariales) form key biogenic habitats in coastal regions of temperate and Arctic seas worldwide, providing ecosystem services valued in the range of billions of dollars annually. Although local evidence suggests that kelp forests are increasingly threatened by a variety of stressors, no comprehensive global analysis of change in kelp abundances currently exists. Here, we build and analyze a global database of kelp time series spanning the past half-century to assess regional and global trends in kelp abundances. We detected a high degree of geographic variation in trends, with regional variability in the direction and magnitude of change far exceeding a small global average decline (instantaneous rate of change = −0.018 y−1). Our analysis identified declines in 38% of ecoregions for which there are data (−0.015 to −0.18 y−1), increases in 27% of ecoregions (0.015 to 0.11 y−1), and no detectable change in 35% of ecoregions. These spatially variable trajectories reflected regional differences in the drivers of change, uncertainty in some regions owing to poor spatial and temporal data coverage, and the dynamic nature of kelp populations. We conclude that although global drivers could be affecting kelp forests at multiple scales, local stressors and regional variation in the effects of these drivers dominate kelp dynamics, in contrast to many other marine and terrestrial foundation species.

Integrating scientific guidance into marine spatial planning.

Marine spatial planning (MSP), whereby areas of the ocean are zoned for different uses, has great potential to reduce or eliminate conflicts between competing management goals, but only if strategically applied. The recent literature overwhelmingly agrees that including stakeholders in these planning processes is critical to success; but, given the countless alternative ways even simple spatial regulations can be configured, how likely is it that a stakeholder-driven process will generate plans that deliver on the promise of MSP? Here, we use a spatially explicit, dynamic bioeconomic model to show that stakeholder-generated plans are doomed to fail in the absence of strong scientific guidance. While strategically placed spatial regulations can improve outcomes remarkably, the vast majority of possible plans fail to achieve this potential. Surprisingly, existing scientific rules of thumb do little to improve outcomes. Here, we develop an alternative approach in which models are used to identify efficient plans, which are then modified by stakeholders. Even if stakeholders alter these initial proposals considerably, results hugely outperform plans guided by scientific rules of thumb. Our results underscore the importance of spatially explicit dynamic models for the management of marine resources and illustrate how such models can be harmoniously integrated into a stakeholder-driven MSP process.

Density derived estimates of standing crop and net primary production in the giant kelp Macrocystis pyrifera

Assemblages of macroalgae are believe to be among the most productive ecosystems in the world, yet difficulties in obtaining direct estimates of biomass and primary production have led to few macroalgal data sets from which the consequences of long-term change can be assessed. We evaluated the validity of using two easily measured population variables (frond density and plant density) to estimate the more difficult to measure variables of standing crop and net primary production (NPP) in the giant kelp Macrocystis pyrifera off southern California. Standing crop was much more strongly correlated to frond density than to plant density. Frond density data collected in summer were particularly useful for estimating annual NPP, explaining nearly 80% of the variation in the NPP from year to year. Data on frond densities also provided a relatively good estimate of seasonal NPP for the season that the data were collected. In contrast, estimates of seasonal and annual NPP derived from plant density data were less reliable. These results indicate that data on frond density collected at the proper time of year can make assessments of NPP by giant kelp more tractable. They also suggest that other easily measured variables that are strongly correlated with standing crop, such as surface canopy area, might serve as similarly useful proxies of NPP.