Although sand deposits are present on many intertidal rocky shores, their effects on species richness, zonation and trophic structure have often been overlooked. This study is the first to recognise sand as an important abiotic factor on South African rocky shores. Rocky shores in the eastern Cape Province of South Africa are subject to extensive sand inundation and are composed of two hard substrata of differing topographies. Four sites on one substratum and six on the other were sampled quantitatively using quadrats. The biota were identified, counted and/ or weighed to provide a matrix of species biomass and numbers in separate zones. This matrix was then analysed using ordination and classification. A total of 321 species were identified which is more than local rocky or sandy shores. While the intermediate disturbance hypothesis would predict high species richness on these shores, it does not fully explain this richness nor the distribution of species assemblages. Habitat heterogeneity, including the dynamics of sand deposits, is strongly influenced by substratum topography and is the most important factor generating species richness. Abrasion by sand (sand scour) causes local reductions in richness but the presence of semi-permanent sand deposits allows habitation by psammophilic and sand-dependent species. As a result the biota of a sand inundated rocky shore includes both a full rocky shore and a large sandy beach component. Substratum topography controls patterns of sand deposition and retention and community analysis showed that samples were clustered primarily according to species richness and secondarily according to substratum type. Ordination of species identified an arc of species assemblages of decreasing levels of sand tolerance. These corresponded to sample groupings so that th assemblages found in various habitats were characterised by particular levels of sand tolerance. The presence of sand has a negative effect on the biomass of primary producers and filter feeders but a positive effect on the biomass of deposit feeders. Because sand is retained to different degrees in different zones, trophic structure varies between zones and to a lesser extent, between rock types. In general, however, the trophic structure of sand inundated rocky shores is similar to that of non-inundated shores.

The relationship between nutrients and community structure is poorly understood in open-coast habitats. I created a system of artificial tidepools, of identical age and physical dimensions, at two sites that differed in wave exposure, and manipulated nutrient levels and the abundance of herbivores. Using these unique field mesocosms, I explored the role of changes in nutrient dynamics and tested two predictive models of community structure in a rocky intertidal community. I modified a simple food-chain model to include the effect of hydrodynamics on nutrient delivery rates and herbivore foraging efficiency. Field experiments demonstrated that nutrients had strong effects on the abundance and productivity of seaweeds. Algal productivity was negatively influenced by herbivory, contrary to model predictions, because species with the potential to increase growth rates when given additional nutrients were virtually eliminated in the presence of herbivores. The effects of both nutrients and herbivory varied in a manner consistent with predicted effects of hydrodynamic forces. Contrary to simple food-chain models, herbivores did not respond to nutrient additions. I assessed nutrient dynamics during low tide, demonstrating that nutrients were rapidly depleted from tidepools. I also examined variation in nutrient uptake rates relative to the experimental treatments described above, for both whole pools and on a biomass-specific basis. Nutrients were almost always removed from pools at the same rate dispensers added them. Uptake rates were significantly correlated with the abundance of fleshy seaweeds. Synthesizing the results of these and other studies, I proposed that the abundance of tidepool seaweeds can be modeled as a function of pool volume, degree of tidal isolation, water flow at high tide, and herbivory. I tested the predictions of a functional group model and evaluated the validity of equating physical and biological disturbances by examining algal diversity and abundance patterns in tidepools along gradients of potential productivity, herbivory, scour and wave exposure. The abundance of functional groups varied along environmental gradients, but not always in a manner consistent with predictions. I suggested that physical and biological processes must be modeled separately, and that better operational definitions of environmental potentials will aid in development of these models. / Graduation date: 1999

Thesis (Ph. D.)--University of Washington, 1993. / Vita. Includes bibliographical references (leaves [145]-169).

Thesis (Ph. D.)--Oregon State University, 1995. / Typescript (photocopy). Includes bibliographical references (leaves 211-237). Also available online.

Thesis (M. Phil.)--University of Hong Kong, 1991.

While community ecologists have traditionally focused on local-scale processes, it has become apparent that a broader perspective, which explores the community-level ramifications of material fluxes within and between ecosystems, is necessary to effectively evaluate bottom-up influences on community structure and dynamics. In this dissertation, I employed ecosystem principles to understand these processes in rocky intertidal communities. I specifically examined the roles of sessile invertebrates in mediating the transfers and transformations of carbon and nitrogen in intertidal ecosystems. First, I quantified the links between nearshore pelagic and rocky intertidal systems. By assimilating suspended particulate organic material (seston), mussels and other sessile invertebrates serve as mediators of material exchange from pelagic to benthic ecosystems. I evaluated these trophic linkages along productivity gradients on the coasts of New Zealand and Oregon, which allowed me to address the influences of seston quality and quantity on the growth and ammonium excretion rates of mussels. My results highlight the necessity of simultaneously considering both seston quantity (total organic particulates) and quality (phytoplankton availability) in evaluating benthic-pelagic coupling. Second, I assessed the utilization of invertebrate-excreted ammonium by macroalgae in high-intertidal pools. Sessile invertebrates not only serve as mediators of material transfer into intertidal ecosystems, they also chemically transform that material, converting particulate organic nitrogen, which is unusable by macroalgae, into ammonium, which algae readily assimilate. l showed that especially in high-zone pools, which are isolated from the ocean for 80% of the time, invertebrate-excreted ammonium is an important nitrogen source for macroalgae. Ammonium accumulated in tide pools and was subsequently taken up by algae. This novel positive interaction influenced community structure: macroalgal species richness increased with the rate of invertebrate-mediated ammonium loading in pools. Finally, by experimentally manipulating macroalgae and invertebrates in laboratory mesocosms, I quantified the effect of ammonium loading on algal growth. I demonstrated that algal nitrogen assimilation rates increased with the rate of ammonium accumulation in tide pools, which resulted in enhanced growth when invertebrates were present. Together, these studies suggest that by merging community and ecosystem perspectives we can gain unique and important insights into the bottom-up processes influencing intertidal systems. / Graduation date: 2004

published_or_final_version / Zoology / Master / Master of Philosophy

A combination of DGPS/GIS mapping, invertebrate surveys and environmental monitoring over one year (July 2004--July 2005) were used to examine the recovery of permanent tidal pools on a salt marsh in the lower Bay of Fundy which underwent an unmanaged restoration after breach of the dyke over 50 years ago. The results are compared to those of a nearby relatively undisturbed reference marsh. / Pools were found to represent a substantial portion of the marsh habitat, comprising 13% of the total marsh area in the recovering marsh and 4.8% in the natural marsh. Observations indicate ice may be an important mechanism of pool formation and growth in these marshes. Water temperature in the pools ranged from freezing (-2°C) to a maximum of 36°C with ice covering pools for up to one month in the winter. Salinity of the pools ranged from near freshwater (4) to hypersaline (41). Environmental variability was mainly driven by climatic conditions with increased tidal flooding of pools at low elevations tending to make the conditions more stable. / A total of 42 macroinvertebrate taxa were identified in pools of the two marshes, with species richness (S) of individual pools ranging from 13 to 23. An estimated 50 years since dyke failure, the invertebrate fauna of pools in the recovering marsh is indistinguishable from that of the reference marsh. No significant differences in macroinvertebrate communities were detected between sampling dates, pool size or pool depth. Non metric multidimentional scaling and hierarchical cluster analysis supported separating the pool invertebrate communities of this study into those occurring in regularly flooded pools, and those occurring in irregularly flooded pools. Average production of pool macroinvertebrates ranged from 1.79 to 4.03 g dry wt m-2, depending on the amount of vegetative cover in the pools. The pools are characterized by low equitability in species abundance and biomass. The numerically dominant organisms of the pools were mites (Acarina), the gastropod Hydrobia tottentei, Tubificidae oligochaetes, and Chironomus sp. larvae as well as copepods and ostracods.

No description available.

Populations of the sympatric intertidal bivalves, Macoma secta and M. nasuta are compared as to reproductive ecology and population dynamics. Histological examination of seasonal gonadal changes indicate that both species undergo dormancy in the winter months and rapid gamete proliferation in the spring. However, the timing of spawning differs. The M. secta population synchronously spawned in August while the M. nasuta population spawned lightly in late spring and more heavily in the fall. Fecundity estimates indicate M. nasuta adds 140% of winter weight in gametes and M. secta only adds 43% of winter weight in gametes. Spat fall was detected in January for both species and was light in intensity, for either species only 24 per m2. Nursery areas were determined for M. nasuta, which corresponds to adult distributions, and for M. secta, which correspond to the upper and lower fringes of the intertidal distribution but not the center. Analysis of variance on distributional data indicate the temporal stability of density and size for M. secta. M. nasuta densities were also temporally stable but sizes were not. Both species demonstrated significant changes in density with tidal height. A measure of seasonal population variability, the Population-Time Mean Square, was found to have zero correlation with tidal height for M. secta, meaning that environmental stress was the same for individuals at each tidal height . This is believed related to the gradient of calm size to tidal height. An argument for density dependence in M. secta is given with emphasis on individual growth and realized population fecundity; the null hypothesis: As density increases, the resources per individual decrease, therefore the growth rate of individuals decreases. Analysis of sympatric populations indicate differing patterns of life history, pointing out that selection has favored these two species coexistence. Analysis of population dynamics indicated that in comparison to M. nasuta (Low Tide Horizon), M. secta (Mid Tide Horizon) was more stable (terminology from Ricketts, Calvin and Hedgpeth, 1969). This supports Castenholz’ (1967) idea that where climates are mild, the intertidal communities are more stable than the subtidal communities.