Weathering The Storm: The Implications Of Wave Exposure On The Distribution, Phenotype And Growth Of A Temperate Reef Fish
Disturbance is a fundamental process that affects the structure and dynamics of populations. Wave action is an important agent of disturbance in coastal marine systems, and the frequency and severity of wave-associated disturbances is forecasted to increase with climate change. Understanding the effects of waves on coastal marine ecosystems, and the ability of organisms to adapt to wave action, is of growing importance. This is particularly true for intertidal/shallow subtidal species that are subjected to varying, sometimes intense, wave action. Most studies to-date have focused on species with limited mobility (e.g., algae and invertebrates), and have used estimates of wave dynamics that are not always relevant to the spatial scales of these organisms and their home ranges. My thesis focuses on the common triplefin, Forsterygion lapillum, an abundant benthic marine fish inhabiting shallow subtidal and intertidal rocky reefs throughout New Zealand. I develop and implement a protocol to characterise wave climates on an ecologically relevant scale. I evaluate the effects of waves on abundance, phenotype, performance, and behaviour of a reef fish. In Chapter 2, I develop and implement a protocol to characterise wave climate at an appropriate scale. The Wellington south coast is exposed to storm waves that develop in the Southern Ocean and propagate up the east coast of New Zealand. I deployed low-cost HOBO acceleration loggers at two depths within each of six locations along the Wellington south coast to record a time series of wave action at twelve sites. Data from my loggers showed substantial spatial and temporal variation in water acceleration due to interactions between waves and local topography. I used a clustering analysis to characterise my 12 sites as either ‘exposed’ or ‘sheltered’. Assignments to these exposure categories did not match with a priori predictions of exposure, suggesting that wave forces experienced by organisms in the shallow subtidal environment may be difficult to assess from surface-based observations of waves. Data were generally well-correlated with an offshore buoy at all sites, and these correlations were stronger for more exposed sites. In Chapter 3, I explored variation in fish density and phenotype through time and as a function of wave exposure. Densities peaked in summer (corresponding to seasonal recruitment) and declined over winter (consistent with increased losses during high-wave periods), and were generally greater at sheltered locations. While body condition was generally highest for fish sampled from exposed sites (consistent with a density-dependent effect on condition and/or enhancement of foraging with increasing water acceleration), other morphological characteristics did not consistently vary with wave exposure. In Chapter 4, I used otoliths to reconstruct of growth histories of individuals to further elucidate the influence of wave exposure on triplefin phenotypes. Recent growth was not influenced by wave exposure, but this was confounded by strong seasonal variation in growth rates. Lifetime growth rate also did not differ with wave exposure, and was strongly influenced by hatch date. I used mixed effects models to appropriately account for the potentially confounding effects of other features on growth, and found that daily growth rates were slightly positively correlated with site-specific daily measures of wave action. This result can potentially account for the elevated body condition of fish at exposed sites (Chapter 3), and it has important implications for fish inhabiting wave exposed coasts. In Chapter 5, I conducted a lab experiment to evaluate feeding ability in relation to simulated wave action. I used fish of a range of sizes, sampled from either a wave-sheltered or a wave-exposed site, and measured their consumption of prey in calm (low flow) conditions, disturbance (high flow) conditions, and immediately following a period of disturbance. Fish consumed fewer prey during disturbance, and more prey during calm conditions (and a similar consumption rate was observed for fish that were assayed after a period of intense wave action). While this pattern held for fish sampled from both populations, fish from wave-exposed sites consumed more prey than fish from sheltered sites, suggesting phenotypic traits (e.g., behavioural or morphological) that shape their feeding efficiency. Collectively my results suggest that organisms that inhabit wave-exposed coastlines may be intimately linked to wave climate. Waves may have direct effects on numbers (reducing densities via induced mortality) and/or indirect effects on the traits, foraging opportunities, and/or body condition of survivors. Species such as the common triplefin may exhibit plasticity in phenotypic traits that enable them to adapt to dynamic and unpredictable environments. Overall, this thesis provides insight into the ability of an intertidal/shallow subtidal species to cope with variable wave action. Such species may exhibit resilience with increasing wave action due to climate change.