Thesis (Ph. D.)--University of Rhode Island. / Bibliography: l. [95]-99.

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Long term data on the North Sea ecosystem are available for phytoplanktonic, zooplanktonic, berithic, fish, and seabird communities. Temporal changes in these have been examined by numerous researchers over the course of the 20th century, their main objective being to determine how the interannual dynamics of these communities are regulated. This study considers the long term ecosystem dynamics, and the mechanisms behind these dynamics, for the North Sea over the latter half of the 20th century, although it focuses upon the time series operated by the Dove Marine Laboratory, and the Continuous Plankton Recorder (CPR) survey in the central-west North Sea region. A review of the literature suggests that long term changes across the North Sea are ultimately driven by two processes. In the northern, western and central areas of the North Sea, the ecosystem is climatically driven, whilst in the southern and eastern regions, the signal of climate is masked by the large arithropogenic nutrient inputs into these regions. A comparison of the Dove and CPR zooplankton time series for the central-west North Sea area found that although their relative year to year fluctuations were similar, large differences were present in the absolute abundances recorded. Model derived catching efficiencies for the two sampling devices suggested that differences in absolute abundances were mainly due in some zooplankton taxa to a greater degree of active avoidance of the CPR sampling device. Further examination of these two series found that the long term zooplankton trends in the central-west North Sea were dissimilar to those observed for other North Sea regions. Inverse relationships between zooplankton abundance, and the position of the Gulf Stream North Wall, and with air temperatures were also observed. These dissimilar trends and inverse relationships were ultimately found to be due to the presence of an internal predation based mechanism. Cmatic influences were also found to indirectly influence the long term dynamics of the benthos in the central-west North Sea The primary factor influencing interannual variation in benthic abundance was phytoplankton productivity (Le. food), which in turn was related to climatic factors. However, at a second central-west North Sea benthic station, situated within a Nephmps norvegicus fishing ground, constant trawling disturbance of the benthos was the primary factor influencing both benthic abundance and species composition. Over the latter half of the 2O1 century, air temperatures and daily sunshine durations have increased in the central-west North Sea region, alongside changes in climatic proxy variables (e.g. NAO index). However, similar trends were not always observed in the biota. Ultimately, the long term dynamics of taxa, communities and ecosystems may be due to direct or indirect factors, yet interactions between a diversiy of internal and external factors, results in the complex behaviour of biological systems over time. This study shows that, a though the centralwest North Sea ecosystem is climatically driven, similar trends between climate and ecosystem components do not necessarily exist, and nor considenng the high comp exity of the ecosystem, should they be expected.

A comprehensive investigation into the relationship between physical forcing and sus- pended particles in the shallow shelf region of the Western English Channel has been conducted, in order to evaluate the temporal dynamics of suspended particle populations. Measurements were taken across tidal cycles and seasons at station L4, part of the Western Channel Observatory (WCO), using the combination of a free-fall microstructure profiler and holographic imaging. Confirmation that L4 is weakly stratified is given, and that the formation of the seasonal thermocline is substantially altered by the spring-neap cy- cle. Stratification is variable and prone to periodic and partial erosion from atmospheric forcing during any point in any season. L4 undergoes moderate turbulent dissipation, principally as a result of tidal forcing. Typically, values of ε do not exceed 10−4 W kg−1 . L4 also exhibits tidal asymmetry, chiefly in response to stratification which, albeit weak, is frequently able to suppress turbulence when generated from the sea bed. The potential energy anomaly is small at L4, as expected for a weakly-stratified environment. Maxi- mum values in summer were shown to not exceed 50 J m−3 . Values of bed stress, τ0 , are rarely greater than around 0.18 N m−2 . Nonetheless, the critical erosion threshold falls below this, and is therefore smaller than that observed in similar locations around the UK. Seasonality in the amount of material resuspended from the seabed is important at L4. The presence of certain biological particles strongly influence particle size and may also determine if a given particle is lifted from the bed. Particles ≥ 200 μm are relatively rare, the site is dominated by particles smaller than this value in line with many other UK sites. Under certain conditions the theoretical maximum limit of particle size, the Kolmogorov length scale, does not hold and many examples of occasions when this threshold is exceeded are shown. This may generate important consequences in subsequent work undertaken at this site and other temperate shelf locations globally, particularly as these results indicate that maximum particle size appears to be governed less by the size of the local turbulent eddies and more by the presence of biological particles. This is another key seasonal component to particle dynamics in the Western English Channel. Phytoplankton populations are readily advected into and out of the L4 site, calling into question the current sampling strategy of the WCO to rely exclusively upon point measurements. Small increases in atmospheric forcing have the ability to rapidly disperse patches of phytoplankton, possibly to the point of cell mortality. Traditional sampling techniques for assessing zooplankton density have been shown to radically underestimate the number of animals present at L4, which will increase error estimates on current ecosystem models.

In this thesis I explore the effect of parameter uncertainty in ocean biogeochemical models on the calculation of carbon uptake by the ocean. The ocean currently absorbs around a quarter of the annual anthropogenic CO2 emissions to the atmosphere [Scholes et al., 2009], slowing the increase in radiative forcing associated with the increasing atmospheric CO2 concentration. Ocean biogeochemical models have been developed to study the role of the ocean ecosystem in this process. Such models consist of a greatly simplified representation of the hugely complex ocean ecosystem. This simplification requires extensive parameterisation of the biological processes that convert inorganic carbon to and from organic carbon in the ocean. The HadOCC ocean biogeochemical model is a Nutrient-Phytoplankton-Zooplankton-Detritus (NPZD) model that is used to represent the role of the ocean ecosystem in the global carbon cycle in the HadCM3 and FAMOUS GCMs. HadOCC uses twenty parameters to control the processes of biological growth, mortality, grazing and detrital sinking that control the uptake and cycling of carbon in the ocean ecosystem. These parameters represent highly complex and in some cases incompletely understood biological processes, and as a result are uncertain in value. A sensitivity analysis is performed to identify the HadOCC parameters that due to uncertainty in value have the greatest possible effect on the exchange of CO2 between the atmosphere and the ocean—the air-sea CO2 flux. These are found to be the parameters that control phytoplankton growth in the well lit surface ocean, the formation of carbonate by marine organisms and the sinking of biological detritus. The uncertainty in these parameters is found to cause changes to the air-sea CO2 flux calculated by the FAMOUS GCM. The initial effect of these changes is equivalent to the order of the error of current estimates of the net annual carbon uptake by the ocean (2.2 ± 0.3 Pg C y−1 [Gruber et al., 2009], 2.2 ± 0.5 Pg C y−1 [Denman et al., 2007]). This indicates that while the effect of ocean biogeochemical parameter uncertainty is non-negligible, it is within the bounds of the uncertainty of the total (inorganic and organic) ocean carbon system, and is considerably less than the uncertainty in the carbon uptake of the terrestrial biosphere [Houghton, 2007]. However, as the ocean plays a crucial role in the global carbon cycle and the regulation of the Earth’s climate, further understanding and better modelling of the role of the ocean ecosystem in the global carbon cycle and its reaction to anthropogenic climate forcing remains important.

Phytoplankter sind die bedeutendsten Primärproduzenten in aquatischen Ökosystemen und leisten ca. 45 % der globalen Primärproduktion. Sie weisen eine hohe taxonomische Diversität mit vielfältigen morphologischen und physiologischen Eigenschaften auf. Funktionelle Eigenschaften, die das licht-, temperatur- und nährstoffabhängige Wachstum sowie Verlustgrößen (Fraß, Sedimentation) beschreiben, sind eine Möglichkeit Phytoplankter zu klassifizieren. Diese Einteilung gewinnt für das Verständnis und die Modellierung der Phytoplanktonsukzession (z.B. in Wassergütemodellen) immer mehr an Bedeutung, da nicht taxonomische, sondern funktionelle Eigenschaften das Vorkommen der Phytoplankter im Gewässer bestimmen. Das Wassergütemodell SALMO ist ein mechanistisches, dynamisches, vertikales 1D-Modell, welches Nährstoffe, Sauerstoff, Detritus, 4 funktionelle Phytoplanktontypen und eine Zooplanktonmischgruppe simuliert. Bei der Modellanwendung wurde ersichtlich, dass die vorhandenen funktionellen Typgruppen teilweise nicht ausreichen, um die funktionelle Diversität der realen Phytoplanktongemeinschaften adäquat widerzuspiegeln. Deshalb wird eine Erweiterung der funktionellen Typgruppen in SALMO angestrebt. Eine empirische Freilanddatenanalyse an zwei morphologisch und trophisch verschiedenen Talsperren ergab insgesamt 10 funktionelle Phytoplanktontypen (Typgruppen 1, 2 und 4 bereits in SALMO vorhanden). Die bestehenden funktionellen Eigenschaften wurden erweitert und teilweise verändert (insgesamt 10 funktionelle Eigenschaften). Die neue funktionelle Typgruppe 6 (unbewegliche, koloniale, gelatinöse Chlorophyceen) wurde in lichtabhängigen Wachstumsversuchen und Fraßversuchen mit Daphnia galeata × hyalina genauer untersucht. Als stellvertretende Art für diese Typgruppe wurde Sphaerocystis schroeteri gewählt. Die Wachstumsversuche bei 20 °C kennzeichneten Sphaerocystis als Starklichtart mit einer relativ hohen Wachstumsrate (Iopt = 250 µmol m-2 s-1; µmax = 0.96 d-1; 12:12 h Hell-Dunkelzyklus). Eigene Nährmediumsversuche wiesen auf eine Abhängigkeit von freiem CO2 hin. Dies bietet eine Erklärung für ihr häufiges Auftreten in oligo- bis mesotrophen Gewässern. Bei Anwesenheit von Daphnia konnte nach 96 h Versuchsdauer eine signifikante Erhöhung der Koloniegröße (Anteil nicht fressbarer Kolonien 3-74 %), im Vergleich zur Kontrolle (Anteil nicht fressbarer Kolonien 0-5 %), festgestellt werden. Dies stellt einen effektiven Fraßschutz gegen größenselektiv filtrierendes Zooplankton wie z.B. Daphnia dar und kann bei hohem Fraßdruck zur Dominanz der Typgruppe führen (z.B. Klarwasserphase in eutrophen Gewässern). Der Effekt war nicht durch Infochemikalien induzierbar. Aus Literaturdaten wurde eine relativ hohes Topt von 27 °C, eine nur mäßige Konkurrentstärke um Phosphor (mittlerer KP-Wert) sowie eine Brutto-Sinkgeschwindigkeit von 1.5 m d-1 entnommen werden. / Phytoplankter are the most important primary producer in aquatic ecosystems and contribute about 45 % to global primary production. Phytoplankton communities usually show an enormous taxonomical diversity. However, besides taxonomical diversity, a phytoplankton can also be characterised by its functional diversity by means of functional traits like resource-dependent growth characteristics (e.g. KI-, KP-, KN-values) or sensitivity to zooplankton grazing and sedimentation. Since functional, and not taxonomic, traits drive the phytoplankton succession, a functional classification of phytoplankton is recognised to be increasingly important for the ecological understanding and modelling of plankton succession (e.g. in water quality models). The mechanistic, dynamic, vertical 1D water quality model SALMO simulates the dynamics of nutrient concentrations (P, N), oxygen, detritus, 4 functional phytoplankton types and 1 mixed zooplankton group. Different model applications showed that the number of phytoplankton types in SALMO was sometimes not sufficient for accounting for the observed functional diversity in phytoplankton communities. Therefore, the aim of this thesis is to extend the functional diversity of phytoplankton types in SALMO. After careful evaluation of empirical data from two reservoirs with different morphology and trophic state 10 functional phytoplankton types are proposed (types 1, 2 and 4 remained unchanged). Existing functional traits in SALMO were extended and partly changed (in total 10 functional traits). For the new functional phytoplankton type 6 (non motile, colonial gelatinous green algae) light-dependent growth rates were measured and grazing experiments with Daphnia galeata × hyalina were carried out in the laboratory. For these experiments, Sphaerocystis schroeteri was chosen as representative species for this group. It is characterised by high light demand and relative high growth rates at 20 °C (µmax = 0.96 d-1, Iopt = 250 µmol m-2 s-1, 12:12 h light-dark cycle). Their growth depends on the availability of free CO2. This explains the common occurrence in oligo-mesotrophic water bodies. The range of the colony sizes of Sphaerocystis increased in the presence of Daphnia significantly (portion of inedible colonies 3-74 %) in contrast to the control (portion of inedible colonies 0-5 %). This was induced by direct size-selective grazing and not by infochemicals. Thus, Sphaerocystis has evolved an effective defence mechanism against size-selective filter-feeding zooplankton and is competitive superior at high zooplankton grazing pressure (e.g. clear water stage in eutrophic waters). The lack of experimentally derived parameters for temperature- and phosphorus-depending growth (Tmin, Topt, KP-value) as well as gross sinking velocity was complemented with literature data. The alga is only moderately competitive for phosphorus and exhibits relatively high minimal and optimal growth temperatures (Tmin = 7 °C, Topt = 27 °C). Since type 6 consists of large, non motile species, gross sinking velocity was estimated to be about 1.5 m d-1.

La Méditerranée Nord-Occidentale (MNO) est marquée par un processus de formation d’eau profonde en hiver qui induit une forte floraison phytoplanctonique au printemps. L'objectif de cette thèse a été de caractériser la dynamique de la communauté mesozooplanctonique à l'échelle régionale de la MNO, y compris dans la zone de convection profonde (ZCP), en évaluant ses stocks, sa composition taxonomique, sa structure en taille et ses liens trophiques avec le phytoplancton, en relation avec l'environnement hydro-biogéochimique. Trois campagnes océanographiques ont été menées en saisons contrastées: hiver, printemps et été, fournissant un jeu de données unique à cette échelle régionale. Le zooplancton est caractérisé par de faibles abondances et biomasses en hiver, surtout dans la ZCP, puis par une augmentation générale printanière, en abondance et en biomasse. Des différences spatiales s’observent, la ZCP présentant les plus forts changements de biomasse de l’hiver au printemps. Les valeurs d'été sont similaires aux valeurs hivernales et sont assez homogènes dans la zone d'étude. L'impact du broutage estimé n'est pas suffisant pour contrôler globalement la floraison printanière. Cependant, au printemps, toute la MNO, à l'exception de la ZCP, subit un contrôle top-down du zooplancton sur le phytoplancton, tandis que dans la ZCP, les valeurs de chlorophylle-a restent élevées malgré la forte demande en carbone du zooplancton, ce qui indique un contrôle bottom-up. Cette étude montre que la ZCP est probablement une zone d'intense transfert d'énergie vers les niveaux trophiques supérieurs ainsi que d'export de matière organique en MNO. / The North-Western Mediterranean Sea (NWMS) is characterized by a deep water convection process in winter, which induces a large phytoplankton bloom. The main objective of this thesis was to characterize the dynamics of the mesozooplankton community at the regional scale of NWMS including the deep convection zone (DCZ), by assessing its stocks, taxonomy and size structure, and by evaluating its phytoplankton-zooplankton trophic links in connection to the hydrological and biogeochemical environment. Three oceanographic cruises were conducted to map the NWMS in contrasting seasons: winter, spring, and summer. This represents a unique data set of zooplankton at this regional scale. The NWMS was characterized in winter by low zooplankton abundance and biomass. In spring, a general increase was found. Spatially DCZ was characterized by lowest stocks in winter and the highest in spring. In summer, biomass and abundance were similar to winter values and were quite homogenous over the study area. The estimated zooplankton grazing impact was not sufficient to globally control the spring phytoplankton bloom. However, in spring, all areas except the DCZ incurred top-down control by zooplankton on the phytoplankton stock. In the DCZ, the chlorophyll-a values remained high despite the high zooplankton biomass and carbon demand, indicating a sustained bottom-up control. This study indicates that the deep convection zone is likely an area of both enhanced energy transfer to higher trophic levels and organic matter export in NWMS.