A comprehensive study involving 52 microtidal beaches spanning from reflective to dissipative states and located in tropical, subtropical and temperate regions in the Atlantic, Pacific and Indian oceans was carried out to unravel the relative roles of latitude and beach morphodynamics in determining beach macrobenthic species richness, abundance, biomass and mean individual body sizes. Since beach slope is one of the most important factors controlling beach fauna, a model based on beach geometry was applied to the sub-aerial beach deposit to understand the role of physical factors in predicting equilibrium beach slopes. Additionally, it was tested if the occurrence of beach types is related to latitude, and how physical factors change with morphodynamics and latitude. The beach morphometric model makes three assumptions: 1) that the cross-section of a beach deposit is equivalent to a right-angled triangle, 2) that the physical hydrodynamic factors (wave height and spring tide amplitude) and the beach deposit characteristics (sand grain size, beach slope and width) are interchangeable with their geometric counterparts producing an equation to explain beach geometry, and 3) that the predicted beach slope is in equilibrium with the hydrodynamic and sedimentological forces. The equation for predicting beach slope was derived and then tested against field data collected over 52 beaches. The predicted slopes were not significantly different from the observed slopes of the studied natural beaches. However some estimated slopes were different than observed ones. Possible sources of deviation between calculated and observed slopes may be the systematic sampling errors associated with field data. Alternatively, observed slopes could be the result of past hydrodynamic conditions, explaining the differences with the slopes calculated by the beach morphometric model, which assumes a state of equilibrium between beach slope and hydrodynamic conditions. A higher correlation of beach faunal structure with observed slopes rather than with present hydrodynamic conditions could then be indicative of faunal responses to previous hydrodynamic conditions in the same way was the observed slopes. The beach morphometric model could therefore be also useful in predicting faunal responses to changing hydrodynamic conditions. Since the model does not consider wave period, it is concluded that further tests should be done using laboratory and time-series field data and incorporating the role of wave period and beach permeability to ascertain its predictive value. Tropical regions had significantly more beaches in a reflective state than temperate and subtropical regions. Some tropical beaches were fronted by coral reefs, which not only provided coarse carbonate particles, but also additionally dissipated the low wave energy present in these climatic areas. Only one dissipative beach with high energy was found in the tropics, in southeast Madagascar. Temperate and subtropical regions, on the other hand, were dominated by dissipative beaches with medium to fine sands. Open oceanic reflective beaches were non-existent in the subtropics and rare in temperate regions, only occurring in estuaries, enclosed bays or on islands facing the continent. Intermediate beaches were more common in the subtropics but also occurred frequently in the other two regions, having higher energy in temperate regions. Reflective beaches had significantly steeper slopes, coarser sands, smaller waves and shorter swash lengths when compared to dissipative beaches. Additionally, reflective beaches were narrower, had deeper water tables and in consequence shorter saturation zones. Wave periods and surfzone widths were on average larger on dissipative than on reflective beaches. The frequency of occurrence of beach types is therefore related to the climatic signature of each latitude. Mid latitudes would be predicted to have more dissipative beaches with large and long waves because of their proximity to the storm generating belt around 50-60° S - these beaches will have a predominantly finer sands because of the input by rivers in rainy areas. Lower latitudes will have more reflective beaches due to a modal low energy wave climate and also because of the presence of inshore and offshore biotic structures such as coral reefs that dissipate even more the energy from the waves. Tropical and subtropical regions had larger marine species pools than temperate regions. After controlling for biogeographical differences in total species pool, dissipative beaches were on average significantly richer than intermediate and reflective beaches. Crustaceans were also more diverse on dissipative beaches, this difference being not significant for either molluscs or polychaetes. Significant relationships were found between total beach species richness (with or without terrestrial species), crustacean and mollusc species richness with beach morphodynamics as represented by the Dean’s index. Indices incorporating the role of tide, such as BSI, were less important in predicting species richness on the microtidal beaches studied here. Several other physical factors were also significantly correlated with species richness, the most important being the Beach Deposit Index, i.e. BDI, a composite index of beach slope and grain size. The highest correlation was between BDI and relative species richness, i.e. local beach species richness / regional species richness. Total regional marine species richness was higher in Madagascar and North Brazil (tropical beaches) than in Southeast Brazil (subtropical), or the West Coast of South Africa and South-Central Chile (temperate beaches). Not only was the pool of species capable of colonizing beaches richer in tropical and subtropical regions, but also local diversity of each morphodynamic beach type was higher among tropical beaches than their temperate counterparts. Although the results of this study agree with the predictions of the swash exclusion hypotheses, several reflective and intermediate beaches had higher species richness than predicted before and this seems to be linked to the presence of finer sediments and a less turbulent flow for larvae to settle. It is concluded that beach species richness is not necessarily controlled by one major morphodynamic parameter; this control is complex and probably involves multiple interacting biotic (biological interactions) and abiotic (e.g. grain size, wave height, beach slope and width, water table) factors. Species richness seems to be controlled on two different scales: on an evolutionary one where tropical and subtropical regions have higher regional and local diversity due to higher speciation rates; and on an ecological scale, where fine grained sand beaches have their carrying capacity enhanced by higher larval settlement rates and survival of recruits towards adulthood. On average dissipative beaches had higher total densities and macrofaunal abundances than reflective ones. Crustaceans, terrestrial species and cirolanid isopods such as Excirolana spp. were also more abundant on dissipative beaches. The difference was not significant for molluscs and polychaetes. Significant relationships were found between total abundance, crustacean species richness and beach morphodynamics as represented by the Dean’s index. Again, BSI was less important than individual physical factors in determining faunal abundance on the studied microtidal beaches. Several other physical factors were also significantly correlated with abundance, the most important being the Beach Deposit Index (BDI). The highest correlation was between BDI and total macrofaunal abundance. Factors related to surf zone processes, and possibly productivity, were highly correlated with total community, crustaceans, Excirolana spp. and terrestrial species abundances. The abundances of polychaetes and molluscs were better correlated with factors related to the beach deposit (BDI, slope, grain size and water table depth). Significant differences were observed between latitudinal regions for the average beach and also for each beach type. In general temperate beaches harboured larger community densities and abundances, and also crustacean, Excirolana spp. and terrestrial abundances. Mollusc and polychaete abundances were larger on subtropical and tropical beaches. The control of abundance on a sandy beach is complex and involves multifactorial processes at evolutionary and ecological scales. At evolutionary scales animals seem to attain higher abundances in the region where they first evolved, e.g. amphipods and isopods in temperate regions. At ecological scales they attain higher abundances where productivity is higher (total macrofaunal, crustaceans, Excirolana spp., terrestrial spp.) or where the settlement environment is more benign (molluscs and polychaetes). Dissipative beaches supported larger average and total community biomass than reflective beaches. Crustaceans, terrestrial species and cirolanid isopods such as Excirolana spp. also had larger biomass on dissipative beaches. The difference was not significant for molluscs and polychaetes. Significant relationships were found between the biomass of community and taxonomic groups with beach morphodynamics as represented by the Dean’s morphodynamic index. On the microtidal beaches studied here, BSI was less important than other morphodynamic indices and single physical factors in determining faunal biomass and mean individual body size. Surf zone characteristics such as wave height, period and surf zone width had the highest correlations with community, crustacean.

The objective of this research was to improve our understanding of how changes in the environment affect ecological processes. Change detection is often confounded by the large variation found in ecological data due to the difficulty of finding replicates in nature. Intertidal communities were chosen for studies of biophysical interactions because the physical gradients are very strong, thus creating complex systems within spatial scales that are easily sampled. The selection of replicate beach habitats was the first step in designing a sampling protocol for comparative analyses of nearshore community structure. A high resolution shoreline partitioning model was developed to quantify the physical attributes of homogeneous shoreline segments and to statistically cluster replicate segments. This model was applied at 3 locations in Washington State. A portion of the south shore of San Juan Island was partitioned and the physical attributes quantified. Three groups of rocky segments differing only in slope angle were selected for biological sampling. The objective was to test the fidelity of macroalgal and invertebrate populations to replicate bedrock shore segments. The results showed that community structure and population abundances were more similar within groups of replicate segments (similar slopes) than among groups (different slopes). In South Puget Sound, community structure was compared to test for a deterministic organization of communities among replicate soft sediment beaches in an estuary. The results showed that replicate beach segments support similar communities, that communities become less similar as the distance between replicates increases, and that replicates within or among nearshore cells with similar temperature and salinity support communities that are more similar than replicates among cells with different water properties regardless of distance. On the outer Olympic coast, community comparisons were made among 9 sand beaches over a shoreline distance of 250 km. The results show that these communities are similar within segments and within nearshore cells, but because of population abundance fluctuations, the communities were different among cells and among years. This study shows that processes determining patterns in nearshore habitats can be quantified, which is a significant contribution to studies of habitat distribution and the siting of marine preserves. / Graduation date: 2000

Seagrasses provide an important habitat for gillfish, crustacea and migratory birds. Extensive losses of seagrass in the Northern Hemisphere have occurred since the 1930's in what has been described as a 'wasting disease'. More recently, point-source contamination by nutrient inflows, herbicides (anti-fouling agents used on commercial shipping), heavy metals and fresh water inflows have helped explain localised losses of seagrass amounting to 20% in the case of Adelaide's metropolitan coastline, South Australia. However, losses of seagrass acreage have also occurred in regions that are far removed from anthropogenic activity and these are less easily explained by pointsource contamination. Intertidal seagrasses, such as Zostera muelleri, are subjected to environmental pressures imposed on them by the marine and terrestrial environments. For the purpose of this thesis, the intertidal environment is regarded as a complex of several components or micro-environments, each imposing a selective pressure or stress upon seagrass. The many stress factors create a tolerance zone in which Z. muelleri can survive. Zostera muelleri has adapted its physiology and biochemistry to the selective pressures that operate within the intertidal region. Zostera muelleri's internal leaf morphology has many gas storage compartments (lacunae) that extend from the leaves to the roots of the plants and its photosynthetic biochemistry has also adapted to the intertidal region enabling the sequestering of carbon under conditions of high irradiance and temperatures. It is evident from the literature that the survival of intertidal seagrasses requires effective photosynthesis. It is also evident that events that interfere with the synthesis, translocation and release of photosynthesised oxygen from the roots of Z. muelleri will compromise seagrass survival. The present study has revealed that herbicides, used in broad- acre farming, can be transported to the intertidal environment and negatively impact upon Z. muelleri. Extensive studies by others have shown that transport mechanisms, such as 'spray drift' and 'run-off', can move herbicides from their point of usage. However, 'dust' (wind-eroded soil ) as a transport mechanism for herbicides to the intertidal environment is less well studied. This is surprising, inasmuch as there is a known rate of pedogenesis in Adelaide of five to ten tonnes per km2 per annum from the accretion of dust. Results of the present study suggest that farmed soils of the Yorke Peninsula have a range of potentials to form fine particulate matter ('dust') and this potential is likely determined by the soil type and farming practices. Soil surface applied herbicides, such as 2,4-D, are 'lost' from land at 5% of the applied rate while soil-incorporated herbicides, such as treflan (trifluralin), are lost at 1.5% of the applied rate. Indeed, such herbicides can be transported as dust for tens to thousands of kilometres. Instrumental analytical techniques used in the present study have detected 2,4-D, trifluralin and sulfonylurea herbicides on whole soil. Additionally, 2,4-D- like chemicals have also been detected in whole soil and in dust obtained from whole soil. Bioassay techniques using Z. muelleri have shown that its photosynthetic pathways are negatively impacted upon by micromolar concentrations of 2,4-D that are similar to the known losses of this herbicide from land. It is concluded that, at these concentrations, 2,4-D acts as an auxin, up-regulating growth in affected plants. Such up-regulation is unlikely to be problematic in terrestrial plants since gas flows to the external environment are largely controlled by stomata. However, seagrasses lack stomata and the auxin-like activity of 2,4-D appears to have a negative impact on Z. muelleri. This is probably caused by an up - regulation in oxygen production and a subsequent oxygen-inhibition of a key enzyme ( ribulose 1,5- bisphosphate carboxylase, RUBISCO ) used in the carbon-sequestering photosynthetic process. The proposed inhibition of RUBISCO is then likely to cause a carbon deficit and a subsequent energy deficit within affected plants. One interpretation of the results presented is that Z. muelleri simply outgrows its intertidal environment after a transient exposure to an auxin-like concentration of 2,4-D. With increasing use of auxin-like herbicides, and the associated increasing stress imposed on photosynthetic processes, it is likely that further negative impacts will occur on intertidal seagrass species. Continued depletion of seagrass acreage will further adversely affect fishing yields unless appropriate measures are not taken. Closer collaboration between regulators, farm managers and herbicide manufacturers is now necessary in order to minimise the negative impact of herbicides on intertidal species. / Thesis (M.App.Sc.)--School of Earth and Environmental Sciences, 2005.

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

The influence of loss of diversity on community dynamics and ecosystem functioning has recently received considerable attention. Although study of biodiversity has a long history within ecology, empirical investigations exploring consequences of loss have been rare. Because many factors confound diversity comparisons, experimental manipulations of diversity offer the most direct way of attributing cause to diversity loss. The effects of reduction in number of species will depend on the strength and sign of species interactions affected by loss of diversity. An experiment performed on a high zone, rocky intertidal community in which macroalgal diversity was manipulated demonstrated that effects of diversity loss will be highly dependent on which species are removed. However, effects of diversity reductions were strongest at the harsh end of a stress gradient where interactions were more positive. Thus, factors that affect the strength and sign of species interactions such as the degree of physical stress may serve as a rough guide to where the effects of diversity loss will be most severe. An assessment of the influence of diversity on community response to a strong physical perturbation was performed using an experimentally-induced thermal stress. Higher diversity treatments were most strongly affected directly by the stress because such treatments had higher abundance and therefore more biomass to lose. However, those same treatments recovered more quickly from the stress. Community recovery of initially low diversity treatments was slowed by persistence of non-typical states or slow recovery of dominant species. A simulation study was performed to assess the ability of different experimental designs to detect biodiversity effects. Our ability to predict consequences of changes in diversity will be dependent on our ability to distinguish the most influential biodiversity "components" within a system. This study uncovered a phenomenon that will be common in biodiversity studies: misidentification of one biodiversity component (e. g., an effect of a keystone species) as a different component (e. g., an effect of the number of species). I call this phenomenon "aliasing." Because of the complexity of biodiversity, experiments and observational studies will be highly susceptible to aliasing and, thus, results will require careful interpretation. / Graduation date: 1997

Thesis (Ph. D.)--Oregon State University, 2002. / Typescript (photocopy). Includes bibliographical references (leaves 229-246). Also available on the World Wide Web.

Thesis (Ph. D.)--University of Washington, 2001. / Vita. Includes bibliographical references (leaves 145-164).

Thesis (Ph. D.)--University of Washington. / Bibliography: l. [144]-150.

Thesis (Ph. D.)--University of Melbourne, 1972. / Includes appendices. Includes bibliographical references (p. 243-263).

Thesis (M.A.)--Central Connecticut State University, 1997. / Thesis advisor: Dr. Kim A. Wilson. " ... in partial fulfillment of the requirements for the degree of Master of Arts in Biology." Includes bibliographical references (leaves 47-56).