published_or_final_version / Biological Sciences / Doctoral / Doctor of Philosophy

Investigations into the supply, settlement and recruitment of the barnacle Semibalanus balanoides (L.) to shores in Fife, East Scotland were undertaken over three consecutive years (2004 – 2006). Several designs of a passive larval trap, based on earlier published designs, were employed to quantify the delivery (supply) of cyprids to the substratum. Pump samples from the water column were collected to provide a measure of intertidal cyprid concentration. Cyprid concentration was found to exhibit both spatial and temporal variation, but was also found to be significantly correlated with cyprid supply, as quantified by the traps. In some years, pump and trap samples suggested that cyprid abundance in the intertidal was mediated by wind-driven processes. An artificial substratum was used to quantify cyprid settlement, allowing investigations into the supply-settlement relationship. Supply and settlement were found to be two very distinct biological phases. Supply describes the flux of larvae to the substratum. Settlement provides a measure of the amount of larvae that decide to settle on the substratum, as initial attachment for barnacle larvae is not permanent. Whilst supply saturation is unfeasible, daily saturation of the substratum by settlers was observed at many sites along the Fife coast. Levels of settlement saturation were found to vary both temporally, between years, and spatially, over scales of km. Varying levels of the desperation of larvae to settle, mediated by dwindling energy reserves, was thought to explain some of this variation. The supply data also provided some evidence of the possible movement of cyprids along the Fife coast from Fife Ness to Tentsmuir. However, wind-induced transport may also play a dominant role on the Fife coast. A comparison of supply and recruitment of larvae into adult populations revealed that both pre- and post-settlement events may influence recruitment. The relative importance of these factors was shown to be density-dependent. Negative intraspecific interactions were only seen in denser aggregations of adults. In less crowded aggregations, recruitment reflected initial patterns of larval supply.

Superior swimming stamina in adult fish is presumably passed on to their offspring, but the ontogeny of the appearance of superior stamina and the requisite enhanced cardio-respiratory support for locomotion in larval fishes has not been determined. Is the expression of the suite of parental traits enabling superior swimming stamina in their offspring dependent upon their achieving juvenile/adult morphology, or does it appear earlier in their larvae? To answer this, adults were classified into three groups based on swimming stamina, followed by measurement of length, mass, and width. Larval offspring from the two parental groups -high stamina larvae (HSL) and low stamina larvae (LSL)- were reared at 27°C in aerated water (21% O2). Routine and active heart rate, routine and active mass specific oxygen consumption were recorded through 21dpf, and cost of transport (COT) and factorial aerobic scope were derived from oxygen consumption measurements. Routine heart rate at 2dpf of LSL was 164 ± 1 b·min-1, compared to only 125 ± 2 b·min-1 for HSL. Routine heart rate subsequently peaked at 203 ± 1 b·min-1 at 5dpf in the HSL group, compared to 207 ± 1 b·min-1, at 4dpf in the LSP larvae. Active heart rate at 5 dpf of LSL was 218 ± 2 b·min-1 compared to 216 ± 2 b·min-1 for HSL. Active heart rate increased slightly to 227 ± 2 b·min-1 for LSL before decreasing again, while active heart rate remained relatively constant for HSL. Routine O2 consumption at 2dpf of HSL was 0.09 μmol·mg-1·hr-1, compared to 0.03 μmol·mg-1·hr-1 in LSL. Routine O2 consumption subsequently peaked at 0.70 μmol·mg-1·hr-1 at 9dpf in the HSL, compared to 0.71 μmol·mg-1·hr-1, at 9dpf in the LSL. These values dramatically decreased before leveling off at around 0.20 μmol·mg-1·hr-1 and 0.15 μmol·mg-1·h-1, respectively. Active O2 consumption at 5dpf for HSL was 0.38 μmol·mg-1·hr-1, compared to 0.57 μmol·mg-1·hr-1 for LSL. Active O2 consumption subsequently peaked at 0.97 μmol·mg-1·hr-1 at 10dpf in HSL, compared to 1.19 μmol·mg-1·hr-1 at 7dpf in LSL. These values also dramatically decreased and leveled off. Significant differences (p < 0.05) in heart rate and oxygen consumption persisted through 21dpf. The onset of differences observed in routine and active heart rate in early larvae, correlated with parent stamina, show that juvenile or adult features are not required as a precondition for the emergence of phenotypic physiological differences.

Freshwater ecosystems are under increasing pressure from a variety of contaminants, including heavy metals from mining operations, which can have complex effects that are difficult to evaluate. To detect early warnings from elevated concentrations of metals, organisms are sought to be used as monitoring tools. For example, mouthpart deformities in Chironomid larvae have been proposed as a bioindicator of stress in aquatic environments. However, the frequency and cause of these deformations, and their sensitivity to different stressors remain uncertain. In this study, I evaluated the usefulness of mouthpart deformities as a tool to monitor the effects of heavy metals from mining in northern Sweden. To do this, the mouthparts of 3789 Chironomid individuals analyzed from 17 sites closely located to mining operations and tested against concentrations of metals and DOC in the water chemistry of lakes and rivers. The frequency of deformities ranged from 0.00 – 4.79 % across all sites. Metal concentrations ranged from ‘very low’ to ‘low’ based on biological effect risk assessments. Of these, copper (R2 = 0.73) and cobalt (R2 = 0.66) were found to be significantly correlated with frequency of deformities. Additionally, the occurrence of deformities declined with DOC concentration, this was a nonlinear relationship. Frequencies of deformities observed in this study were lower than what have been reported to similar studies. The result from this study, together with other studies, suggest that deformities in Chironomid larvae are sensitive to even low levels of certain metals and could potentially be a good biomonitoring tool for early warnings of contamination in freshwater environments.

Les huîtres présentent un cycle de vie en deux phases : les larves pélagiques s’adhérent avant de se métamorphoser pour une vie benthique.L’adhésion larvaire se fait au stade pédivéligère par sécrétion d’un bioadhésif produit par un organe spécialisé : le pied. Bien que l’huître Crassostrea gigas soit un organisme d’importance économique et écologique, et un modèle d’étude en biologie marine, le phénomène d’adhésion chez la larve pédivéligère est peu documenté. Une étude morphologique des larves pédivéligères par histologie et microscopie électronique a été réalisée, afin de décrire les glandes responsables de la sécrétion de l’adhésif. Une composition majoritairement protéique de l’adhésif a été révélée par histochimie et spectroscopie FTIR.Une analyse in silico des données transcriptomiques disponibles chez C. gigas a permis d’identifier des gènes probablement impliqués dans l’adhésion.Deux analyses protéomiques, menées sur les larves entières et l’adhésif sécrété ont permis de caractériser des protéines en lien avec la biosynthèse et la structure de l’adhésif. Une protéine de type collagène apparaît impliquée dans la structure de l’adhésif de C. gigas. Cette première approche de l’étude de l’adhésion de C. gigas, permet d’envisager la valorisation biotechnologique des molécules identifiées. Le développement d’adhésifs biomimétiques, élaborés sur le principe des bioadhésifs marins, autoriserait le collage en milieu humide, et serait une alternative aux adhésifs synthétiques qui malgré leur toxicité, dominent le marché mondial. / Oysters show a two-phase life cycle: pelagic larvae adhere before metamorphosis into benthic life. Larval adhesion occurs at the pediveliger stage by secretion of a bioadhesive produced by a specialized organ: the foot. The oyster Crassostrea gigas is an organism of economic and ecological importance, and a model for study in marine biology, but the phenomenon of adhesion in the pediveliger larvae is poorly documented. A morphological description of the pediveliger larvae by histology and electron microscopy was performed to describe the glands responsible for the secretion of the adhesive.A predominantly proteinaceous composition of the adhesive was revealed by histochemistry and FTIR spectroscopy. An in silico analysis of available transcriptomic data from C. gigas was made to identify genes probably involved in adhesion. Two proteomic analyses, performed on whole larvae and on the secreted adhesive, characterizing proteins related to biosynthesis and adhesive structure. A collagen-like protein appears to be involved in the adhesive structure of C. gigas. This first approach to the study of the adhesion of C. gigas makes it possible to consider the biotechnological enhancement of the identified molecules. Despite their toxicity, synthetic adhesives dominate the world market. The development of biomimetic adhesives, based on marine bioadhesive strategies could be an alternative, and allowing furthermore bonding in wet condition.

O principal objetivo desse trabalho foi analisar a influência das condições ambientais na higidez das larvas de engraulidídeos da plataforma continental ao largo de Santos (24o49\'S - 23o49\'S e 46o55\'W - 45o24\'W). As coletas foram realizadas em dois cruzeiros oceanográficos (setembro de 2005 e março de 2006). Verificou-se diferença nas condições hidrográficas entre as duas campanhas de coletas, sendo que em setembro de 2005 houve maior influência de descarga de águas estuarinas na plataforma, enquanto em março de 2006 houve intrusão da Água Central do Atlântico Sul. As duas situações geraram estratificação vertical e favoreceram a entrada de nutrientes no sistema, o que gera enriquecimento da zona eufótica e beneficia as cadeias tróficas locais. Para avaliar a condição de higidez das larvas, foram utilizados indicadores morfológicos (relação massa-comprimento, fator de condição, altura do corpo da larva, relação entre altura da cabeça e diâmetro do olho) e bioquímicos (quantidade de proteína e relação proteína/DNA). Os indicadores apontaram melhor condição relativa, para as larvas coletadas em março de 2006, que pode ser conseqüência da ocorrência de uma intensa estratificação vertical na coluna de água e de uma maior estabilidade gravitacional em março de 2006, proporcionando maior concentração de nutrientes e ocasionando aumento na densidade e melhora na condição das populações planctônicas. / The aim of this study was to analyze influence of environmental conditions on health of engraulidid larvae of the continental shelf off Santos (24º49´S - 23º49´S e 46º55´W - 45º24´W). Samples were collected during two oceanographic cruises (September 2005 and March 2006). Differences among hydrographic conditions were verified between surveys: in September there was a major influence of the estuarine water discharge in the inner shelf, whereas in March there was an intrusion of the South Atlantic Central Water (SACW) in the region. The two conditions generated vertical stratification and improved the pelagic food web, in which fish larvae participate. Morphological (weight-length relation, condition factor, larvae body height, head height-eye diameter relation) and biochemical (protein quantity and protein/DNA relation) indicators were used to evaluate the larvae health condition. Results showed better relative condition of larvae collected in March 2006 that can be a consequence of the intense water column vertical stratification and gravitational stability in March 2006, providing higher nutrients concentration, increasing density of organisms and improving condition of planktonic populations.

xx, 198 p. A print copy of this thesis is available through the UO Libraries. Search the library catalog for the location and call number. / "Bathymodiolus" childressi is a mixotrophic mussel from Gulf of Mexico cold seeps. There is no genetic differentiation of mussels among the seeps, suggesting wide dispersal of their larvae. This dissertation describes larval biology, ecology, and recruitment dynamics for "B." childressi. Cleavage is spiral at a rate of one per 3-9 hours, with blastula larvae hatching by 40 hours at 7-8 à à °C. At 12-14 à à °C, D-shell veligers developed by day 8 without being fed. Egg size and shell morphology indicate planktotrophy, but feeding was not observed. Embryos developed normally from 7-15 à à °C and 35-45 ppt. Although survival of larvae declined with temperature, some survived at 25 à à °C. Larval survivorship was similar at 35 and 45 ppt. Oxygen consumption increased from blastulae to trochophores and was higher for "B." childressi than for shallow-water mussel trochophores. Estimated energy content of "B." childressi eggs was greater than the energy content of shallow-water mussel eggs. An energetic model predicts that the eggs provide sufficient energy for "B." childressi trochophores to migrate into the euphotic zone. In fact, "B." childressi veligers were found in plankton tows of surface waters. The influence of recruitment on fine-scale distributions of adults at the Brine Pool cold seep was examined through manipulative field experiments. The "Bathymodiolus" childressi population at this site has a distinct bimodal size structure that shifts across an environmental gradient. New recruits of "B." childressi are abundant in the inner zone, where methane and oxygen are high and sulfide is low, leading to the inference that larvae settle preferentially there. Experiments were placed in the inner and outer zones and 2-m away from the bed. The number of larvae collected in traps did not differ among the three zones, nor did settlement density. Juveniles survived and grew in all zones, but more caged than uncaged juveniles survived. Mortality of uncaged juveniles was similar in all zones, suggesting that predation does not cause the bimodal distribution. These results suggest that the bi-modal distribution cannot be attributed to settlement preferences or juvenile mortality, but instead to migration or early post-settlement mortality. This dissertation includes my co-authored materials. / Adviser: Craig M. Young

A Baía do Almirantado (Ilha Rei George, Antártica) possui diversas pequenas enseadas de águas rasas, que sustentam uma abundante fauna bentônica. O regime de marés e de ventos é responsável pelo hidrodinamismo na baía e pela interação com as águas do Estreito de Bransfield. Portanto, trata-se de uma área que pode apresentar uma elevada abundância de larvas de invertebrados marinhos. A distribuição espaço-temporal de larvas pelágicas em águas rasas costeiras da Baía do Almirantado foi estudada entre outubro de 2002 e maio de 2003, e outubro de 2003 e agosto de 2004 em 12 estações rasas (0 a 15 m, ou até 30 m). Também foram amostradas, sazonalmente, quatro estações de maior profundidade, entre 0 a 30 m e 30 a 130 m da coluna de água, entre outubro de 2003 e abril de 2004. Amostras de plâncton foram coletadas com rede cônica de plâncton com malha de 150 &#181;m nas três enseadas da baía (Martel, Mackellar e Ezcurra) e na entrada do seu canal principal. Larvas de diferentes grupos ocorreram em todo o período de estudo. No primeiro ano, a abundância total foi maior que no segundo e, em especial, no verão. As larvas véliger de Gastropoda foram mais abundantes, representando 84% do total do meroplâncton, no primeiro ano de amostragem. No ano seguinte, larvas trocófora foram predominantes (31% do total), seguidas de Gastropoda e Nemertea. Os resultados mostraram uma evidente variação interanual da densidade e da composição do meroplâncton, mas sem um padrão nítido de sazonalidade para determinados grupos. O congelamento da baía no inverno do primeiro ano de amostragem (2002) pode ter afetado a reprodução dos invertebrados bentônicos e a produção de larvas. Quanto à distribuição batimétrica, a abundância média do meroplâncton total foi maior na camada profunda do que na de superfície, na primavera-verão, mas espacialmente a densidade foi semelhante entre as áreas nas distintas camadas. A variabilidade na abundância e composição larval pode estar relacionada à hidrografia e a padrões reprodutivos das espécies bentônicas com desenvolvimento indireto, cujos ciclos reprodutivos são afetados por características ambientais. Em termos gerais, o meroplâncton coletado na Baía do Almirantado foi qualitativamente similar (quanto aos grupos de larvas) ao descrito para outras partes da Antártica. / Admiralty Bay (King George Island, Antarctica) has many small inlets of shallow waters, which supports an abundant benthic fauna. Tidal and wind regime are responsible for the hydrodynamics in the bay and for the interaction with the waters of the Bransfield Strait. Therefore, it is an area that can present a high abundance of marine invertebrate larvae. Spatio-temporal distribution of pelagic larvae in shallow coastal waters of Admiralty Bay was studied between October of 2002 and May of 2003, and October of 2003 and August of 2004 in twelve shallow stations (0 to 15 m, or up to 30 m). Four deeper stations were also seasonally sampled (0 to 30 m and 30 to 130 m of water column) between October of 2003 and April of 2004. Plankton samples were collected with a conical net, with mesh size of 150 &#181;m, in the three inlets of the bay (Martel, Mackellar and Ezcurra) and in the entrance of its main channel. Larvae of different groups occurred in the whole studied period. In the first year, the total abundance was higher than in the second and, especially, in the summer. The veliger larvae of Gastropoda were more abundant, representing 84% of the total meroplankton in the first year of sampling. In the following year, trochophore larvae were predominant (31% of the total), followed by Gastropoda and Nemertea. The results showed an evident interannual composition and density variation of meroplankton, but without a clear pattern of seasonality for certain groups. The freezing of the bay in the winter of the first year could have influenced the benthic invertebrate\'s reproduction and larvae production. Regarding to the bathymetric distribution, the mean abundance of the total meroplankton was higher in the deeper layer than in the surface in spring-summer, however, spatially the density was similar among the areas in the different layers. The variability in the larval composition and abundance may be related to the local hydrography and specific reproductive pattern of the benthic species with indirect development, whose reproductive cycles are affected by environmental characteristics. In general terms, the meroplankton collected in Admiralty Bay was qualitatively similar (in view of the larval groups) to the described for other parts of Antarctica.

O presente projeto analisou o perfil dos ácidos graxos, embriogênese e ontogenia do sistema digestório das larvas da garoupa verdadeira Epinephelus marginatus durante os primeiros dias de desenvolvimento. Além disso, descrevemos como os embriões se dividem, eclodem e desenvolvem suas principais características, e como se desenvolve seu sistema digestório ao longo dos primeiros dias de desenvolvimento, e como utilizam os AG durante os primeiros 8 dias de desenvolvimento. Observamos que as larvas da garoupa apresentam desenvolvimento de suas estruturas digestórias relativamente lento, no entanto, estas são capazes de capturar, ingerir e digerir presas já a partir do 4º DAE. Os ovos da garoupa verdadeira são compostos por elevados percentuais de PUFA nos fosfolipídios e para o processo de eclosão utilizam preferencialmente os SFA dos fosfolipídios. Além disso, os PUFA da série n3 sobrepõe-se aos da série n6, principalmente o DHA, que apresentaram valores elevados em comparação com outras espécies marinhas tanto nos fosfolipídios quanto nos triglicérides nos três primeiros dias de desenvolvimento. As larvas apresentam uma elevada necessidade dos HUFAs DHA/EPA, e durante essa fase é importante a utilização de alimento vivo de tamanho reduzido (copépodes ou &ldquo;SS strain&rdquo; Brachionus rotundiformes) enriquecidos com valores da relação entre DHA/EPA acima de 2,0. Com isso, todo o conhecimento gerado durante esses dias de desenvolvimento já permitem a aplicação deste conhecimento no processo de embriogênese e larvicultura da garoupa verdadeira contribuindo para alavancar sua domesticação e produção em cativeiro e elaborar futuramente um programa de repovoamento desta espécie contribuindo para sua conservação. / This project analyzed the fatty acids profile, embryogenesis and ontogeny of the digestive system of dusky grouper Epinephelus marginatus larvae during the first days of development. Furthermore, we described how the embryos are divided, hatch and develop its main features, and how develops its digestive system during the first days of development, and how they use the FA during the first 8 days of development. We observed that the grouper larvae present relative slow development of their digestive structures, however, the larvae are able to capture, ingest and digest preys already from 4º DAE. The dusky grouper eggs are composed by high percentages of PUFA in phospholipids and for the hatching process it uses preferably the SFA of phospholipids. Additionally, the n3 series PUFAs overlaps the n6 series, especially DHA, which exhibited high values compared to other marine species both on phospholipids as in the triglycerides during the first three days of development. The larvae exhibit a high requirement of HUFAs DHA/EPA, and during this phase is important to use live food of small size (copepods or &ldquo;SS strain&rdquo; Brachionus rotundiformes) enriched with the ratio of DHA/EPA levels above 2.0. Thus, all the knowledge generated during these days of development allow us the application of this knowledge in the embryogenesis and hatchery process of the dusky grouper contributing to leverage its domestication and production in captivity and eventually draw up a restocking program of this species contributing to their conservation.

A primary goal of marine ecology is to understand the physical and biological mechanisms that drive patterns of larval dispersal and population connectivity. The larvae of many marine organisms develop in the pelagic environment before settling on benthic habitat. Early efforts to predict dispersal patterns assumed that larvae are passive particles dispersed by currents for 10s to 100s of kilometers (km). However, recent studies using parentage analyses to observe dispersal patterns reveal that peak dispersal occurs within 1 – 2 km, and declines exponentially with increasing distance from the natal reef. This gap between predicted and observed dispersal patterns suggests that other factors, such as larval behavior, may play an important role in shaping patterns of dispersal. However, due to challenges associated with obtaining larvae early in development, there is little information on the ontogeny of behavior from hatching through settlement. In this dissertation, I begin to address these challenges by developing a protocol for rearing the larvae of two species of sponge-dwelling neon gobies, Elacatinus lori and E. colini. Using lab-reared specimens, I provide the first description of larval development for both species. Then, I investigate the ontogeny of swimming ability in larvae of E. lori, E. colini and the model species Amphiprion percula. Remarkably, A. percula were capable of swimming twice as fast and three orders of magnitude longer than E. lori near settlement. Relating swimming speed to published dispersal patterns for E. lori, A. percula and another species Plectropomus leopardus, I show there is a positive association between swimming speed and the median and maximum dispersal distance. This finding suggests that swimming abilities may influence the extent of long distance dispersal. Finally, I investigate the role of habitat preferences and post-settlement persistence in establishing the distribution of E. lori settlers on sponge habitat. I demonstrate that E. lori settlers are more abundant, persist longer, and prefer to settle on large Aplysina fistularis, suggesting that settling E. lori choose sponge habitats that confer the highest relative fitness. This dissertation indicates the importance of larval and settler behaviors in determining the dispersal patterns and distribution of fishes on coral reefs.