Thesis (M.S.)--University of North Carolina Wilmington, 2009. / Title from PDF title page (February 21, 2010) Includes bibliographical references (p. 35-39)

Coastal transition zones form important interlinking regions where marine ecosystems, rivers and estuarine environments significantly influence each other. Coastal rocky shores are key habitats that sustain a variety of primary producers and invertebrates and due to the dynamic nature of coastal ecosystems, suspension-feeders on rocky shores can be influenced by an array of autochthonous and allochthonous food sources. Fatty acid and stable isotope trophic markers were employed to distinguish between regional and temporal changes in the potential food sources to rocky shore suspension-feeders in the Eastern Cape Province of South Africa. The primary aim was to assess the spatial and temporal influences of contrasting river flows on the available food sources to three indigenous coastal suspensionfeeders, namely the volcano barnacle Tetraclita serrata, brown mussel Perna perna and tubebuilding polychaete Gunnarea capensis. This was done by examining the intra- and interspecific changes in the fatty acid and stable isotope signatures of the barnacles, mussels and polychaetes in the adjacent marine environment of a freshwater-restricted (Kariega) and freshwater-dominated (Great Fish) estuary during austral summer and winter. Multivariate and Kruskal-Wallis analyses of variance on the fatty acid and isotopic signatures, respectively, identified significant regional changes in the barnacles and mussels, while only stable isotopes distinguished between the Kariega and Great Fish polychaetes (P < 0.05). In addition, significant temporal changes were observed in consumer fatty acids and isotope values in both regions (P < 0.05). Bacterial sources, detritus and phytoplankton assemblages, which are influenced by hydrology and vegetation, differed between regions and were mainly responsible for the regional and temporal separations. Principal component analyses on the consumer fatty acid signatures distinguished between animals situated upstream (i.e. north) and downstream (i.e. south) of the Kariega Estuary mouth in summer. The north/south separation was mainly due to greater contributions of diatoms to northernlocated animals and dinoflagellates and detritus to southern-located consumers. In addition, the south-flowing Agulhas Current on the eastern shores of southern Africa appeared to influence the north/south separation in the Kariega region, as water leaving the estuary was probably entrained into the south-easterly flowing currents, thereby depositing estuarinederived detritus to southern populations. In general, diatoms and detritus were essential food sources to the filter-feeders in summer, and flagellates, diatoms, zooplankton and detritus were important in winter. Coastal macroalgae was a key food source in the Kariega and Great Fish regions during both seasons. Consistently large levels of diatom markers (16:1n-7 and 20:5n-3) and dinoflagellate markers (22:6n-3) in consumer tissues in the Kariega and Great Fish regions identified that phytoplankton was their dominant food source. The barnacles, mussels and polychaetes had similar fatty acid markers and a fairly narrow δ¹³C range (-16.5 to -14.4 ‰), suggesting that they probably consumed similar food. Carbon isotope analyses, however, separated the suspension-feeders into slightly depleted (barnacles; -16.5 to -16.1 ‰), intermediate (mussels; -15.8 to -15.0 ‰) and enriched (polychaetes; -15.0 to -14.4 ‰) consumers, but did not provide conclusive evidence of their preferences for specific phytoplankton. Conversely, fatty acid analyses highlighted that barnacles and mussels had greater proportions of dinoflagellate markers (22:6n-3; 7.0-15.3 % TFA), while polychaetes had larger diatom levels (20:5n-3; 15.1-22.2 % TFA). In addition, all three species had consistently large contributions from bacterial fatty acids (15:0, i-16:0, 17:0 and i-18:0; 4.2-13.6 % TFA) in summer and winter, and large proportions of saturated fatty acids (33.3-53.1 % TFA) including those with 14 to 18 carbons, indicating that bacterial and detritus food sources played an important role in their diets. Barnacles had small levels of terrestrial markers (18:2n-6 and 18:3n-3; <2.5 % TFA) and demonstrated increased omnivorous feeding compared with the other suspension-feeders [increased levels of 20:1n-11 and 20:1n-9, higher 18:1n-9/18:1n-7 ratios at ~2.1, enriched δ¹⁵N values at ~10.6 ‰; zooplankton (potentially including microzooplankton, larvae and protists) contribution of up to 61 % of the diet]. Mussels contained significant proportions of the terrestrial markers (18:2n-6 and 18:3n-3; >2.5 % TFA) and exhibited intermediate omnivory (intermediate levels of 20:1n-11 and 20:1n-9, intermediate 18:1n-9/18:1n-7 ratios at ~1.3, less enriched δ¹⁵N values at ~7.9 ‰; zooplankton contribution of 10-15 % of the diet). The more depleted nitrogen signatures in the mussels relative to the barnacles and polychaetes possibly illustrated a stronger preference for autotrophic food. Polychaetes mainly consumed plant food sources (i.e. microalgae, macroalgae and detritus; high levels of i-18:0, 18:1n-9, 18:4n-3 and 20:5n-3) and displayed little omnivory (low levels of 20:1n-11 and 20:1n-9, low 18:1n-9/18:1n-7 ratios at ~0.4, intermediate δ¹⁵N values at ~9.1 ‰; zooplankton contribution of <10 % of the diet). The barnacles, mussels and polychaetes are all suspension-feeders, originally presumed to consume the same food sources. The variations observed among the species, therefore, may result from differences in the proportional contributions of the various food sources to their diets as well as distinctions in metabolism. The distinct changes in the fatty acid and stable isotope signatures in all three filter-feeders in the Kariega and Great Fish regions are likely influenced by the diversity in regional vegetation and hydrology in the different systems, combined with interspecific differences in resource partitioning among the species.

Aus Patagonien lagen bisher wenige Daten zu paläoökologischen Veränderungen und zur Paläobiodiversität an der Kreide-Paläogen (K/Pg)-Grenze vor. Um diese Lücke zu schließen, wurden in den argentinischen Provinzen Neuquén und Chubut zahlreiche Profile im Bereich der K/Pg-Grenze detailliert beprobt. Während einer Planktonkrise, wie sie für die K/Pg-Grenze angenommen wird, werden für benthische Mollusken folgende Entwicklungen erwartet: 1. eine Abnahme der Individuenzahlen der Benthosorganismen, 2. eine Abnahme der vom Plankton als Nahrungsquelle direkt abhängigen Organismen, 3. eine Abnahme der Benthonten mit planktotrophen Larvalstadien, 4. eine Abnahme der mittleren Körpergröße der Individuen benthischer Faunengemeinschaften und 5. eine Abnahme der Organismen mit hoher Stoffwechselrate bzw. aktiver, mobiler Lebensweise. Tatsächlich waren die hungerresistenten Gruppen der Nuculoida (flach-infaunale Depositfresser) und der Lucinidae (Chemosymbionten) im Danium signifikant häufiger als im Maastrichtium, während der Faunenanteil der Suspensionsfresser im Danium im Vergleich zum Maastrichtium signifikant niedriger ist. In den Faunengemeinschaften des Daniums von Bajo de Añelo konnten Abnahmen der Individuenzahlen, der mittleren Mobilität und der mittleren Größe festgestellt werden. Die beobachteten paläoökologischen Veränderungen zeigen somit eine Selektivität, die sich auch in der Paläobiodiversitätsdynamik widerspiegelt. Auch hier sind Suspensionsfresser, die direkt vom Planktonangebot abhängig sind, stärker betroffen als Chemosymbionten und flach-infaunale Depositfresser. Die Aussterberaten für Bivalven sind mit 28% moderat. Insgesamt sind die festgestellten Selektivitätsmuster konsistent mit den zu erwartenden Folgen einer Planktonkrise. Trotz einiger regionaler Unterschiede handelte es sich um ein globales Ereignis an der K/Pg-Grenze, das Nord- und Südhalbkugel gleichermaßen betroffen hat. / From Patagonia only a few data on palaeoecological changes and on palaeobiodiversity at the Cretaceous/Paleogene (K/Pg)-boundary are available so far. Therefore detailed samples in different K/Pg-sections in the Argentinian provinces of Neuquén and Chubut were taken to increase the data set for this region. The following trends are expected from a crisis of primary productivity in the aftermath of the K/Pg-boundary: 1. a reduction in the number of individuals of benthic organisms, 2. a decrease in the relative abundance of organisms which rely directly on photosynthesis, 3. a lower proportion of individuals with planktotrophic larval stages, 4. a reduction in the average body size of individuals within palaeocommunities and 5. an increased abundance of organisms with low metabolic rates or inactive lifestyles. In fact the proportion of the starvation-resistant groups of the Nuculoida (shallow-infaunal deposit feeders) and Lucinidae (chemosymbionts) were significantly higher in the Danian compared to the Maastrichtian. In contrast to this, the proportion of suspension feeders in the palaeocommunities was significantly lower in the Danian in comparison with the Maastrichtian. In the Danian benthic assemblages of the Bajo de Añelo area, a decrease in number of individuals, mean mobility and mean size was recognised after the K/Pg-boundary. Thus the palaeoecological changes display a selectivity, which is also reflected in the palaeobiodiversity. Suspension feeders, which depend directly on plankton particles from the water column, show higher extinction rates than chemosymbionts and shallow-infaunal deposit feeders. The extinction rates of Patagonian bivalves are moderate (28%). Overall, the recognized selectivity patterns are consistent with the expected aftermath of a crisis in primary productivity. Despite some local differences, the results argue for a global event at the K/Pg-boundary during which the northern and southern hemispheres were similarly affected.

Numerous ecological studies have used lipids to determine trophic pathways in aquatic systems, as fatty acid profiles provide time-integrated information on an organism’s assimilated diet. Many of these studies have, however, been based on sample collections with a limited temporal scale. The trophic ecology of pelagic systems has been studied intensively using fatty acid analyses, but very little work has been directed toward benthic communities, with the intertidal being especially neglected. The investigation of trophic pathways within rocky shore communities will help us to better understand system responses to environmental changes. The determination of long term temporal variation of the food web within a community could reveal the type, magnitude, duration and frequency of highly seasonal productivity. Changes in fatty acid profiles through time in primary consumers of intertidal rocky shores are poorly understood, but represent an important step towards a more comprehensive understanding of rocky shore food webs, compared with those derived from snapshot or short-term studies. The aim of this thesis was to clarify the temporal variability in the diets of rocky shore intertidal suspension-feeders (the brown mussels Perna perna and the Cape reef worm, Gunnarea gaimardi) and grazers (the Cape sea urchin Parencinus angulosus and the Goat-eye limpet, Cymbulus oculus) on the south east coast of South Africa using fatty acid profiles, and to investigate the effects of life style (e.g. feeding mode) and life cycle on temporal variations in tissue fatty acid profiles. I had three hypotheses: firstly, that suspension-feeders experience high levels of variability in their diets through time because water quality has the potential to change quickly and drastically, whereas grazers experience less variability in their diets over time since their food sources are more constant. Secondly, the reproductive cycles of the suspension-feeder P. perna and the grazer P. angulosus affect the fatty acid composition of their gonads, with temporal variations in lipid composition reflecting changes in reproduction investment. Thirdly, the total amount of energetic reserves available for reproduction are different for each gender (females allocate more energy to egg production than males allocate to gamete production). To address these aims, fatty acid profiles of suspension-feeders and grazers were investigated over a period of twelve months (from July 2010 to June 2011) at a single site on the south east coast of South Africa. The results showed high variability in the fatty acid composition of both the suspension-feeders strongly related with changes in their food source (suspended particulate material). Furthermore, similar temporal changes in fatty acid profiles of the two suspension-feeders were observed over time, reflecting their common diet and life style. There were some inter-specific differences in the suspension-feeders, likely originating from differences in their particle capturing mechanisms. Grazers showed less variability through time compared with the suspension-feeders, with the limpets being more consistent than the sea urchins. The temporal variability in the sea urchin diets may have resulted from the highly diverse and heterogeneous food sources available to them, whereas limpets may be more selective and have a limited range of diet items. Differences between the two grazer species may have arose from differences in their feeding strategies and intertidal zonation. The fatty acid compositions of gonad tissues in both P. perna and P. angulosus showed temporal variability strongly related to reproductive cycle. Differences in the fatty acid values between females and males were apparent, with females richer in total and polyunsaturated fatty acids than males. Spawning and gametogenesis influenced the variability of fatty acids through time in both species, suggesting the importance of considering the reproductive cycle when studying lipids in rocky shore species. Little evidence of lipid transfer between muscles and gonads was seen, suggesting the importance of direct lipid storage into the reproductive tissues. The influence of diet and life history of intertidal consumers on the temporal variability of their fatty acid compositions is important to understand, as it provides us with a better understanding of the functioning of rocky shore systems. There is an enormous potential for future research in this field of study.