Freshwater cyclopoid copepods exhibit at least a fivefold range in somatic genome size and a mechanism, chromatin diminution, which could account for much of this interspecific variation. These attributes suggest that copepods are well suited to studies of genome size evolution. We tested the nucleotypic hypothesis of genome size evolution, which poses that variation in genome size is adaptive due to the 'bulk' effects of both coding and noncoding DNA on cell size and division rates, and their correlates. We found a significant inverse correlation between genome size and developmental (growth) rate in five freshwater cyclopoid species at three temperatures. That is, species with smaller genomes developed faster. Species with smaller genomes had significantly smaller bodies at 22°C, but not at cooler and warmer temperatures. Species with smaller genomes developed faster at all three temperatures, but had smaller bodies only at 22°C. We propose a model of life history evolution that adds genome size and cell cycle dynamics to the suite of characters on which selection may act to mold life histories and to influence the distribution of traits among different habitats.

Copepods in high elevation lakes and ponds in Colorado are exposed to significant levels of ultraviolet radiation (UV), necessitating development of UV avoidance behavior and photoprotective physiological adaptations. The copepods are brightly pigmented due to accumulation of astaxanthin, a carotenoid which has photoprotective and antioxidant properties. Astaxanthin interacts with a crustacyanin-like protein, shifting its absorbance from 473 nm (hydrophobic free form, appears red) to 632 nm (protein-bound complex, appears blue). In six sites in Colorado, habitat-specific coloration patterns related to carotenoprotein complex have been observed. The objective of this study was to determine whether pigment accumulation or carotenoprotein expression has a greater effect on resistance to UV exposure. For each site, copepod tolerance to UV was assessed by survivorship during UV exposure trials. Average UV exposure was determined for each habitat. Astaxanthin profiles were generated for copepods in each site. Ability to withstand UV exposure during exposure trials was significantly different between color morphs (p < 0.0001). Red copepods were found to tolerate 2-fold greater levels of UVB than blue or mixed copepods. Additionally, red copepods have much higher levels of total astaxanthin than blue or mixed copepods (p < 0.0001) and receive a higher daily UV dose (p < 0.0003). Diaptomid carotenoprotein sequence is not homologous with that of other crustaceans in which crustacyanin has been characterized which prevented quantification of carotenoprotein transcript expression. Overall, diaptomid color morph may be an important indicator of UV conditions in high elevation lentic ecosystems.

Copepods comprise an ecologically important role in freshwater and marine ecosystems, which is why they are often considered an important ecotoxicological model organism. The International Organization for Standardization’s (ISO) 14669 protocol is the only guideline for the determination of acute toxicity in three European marine copepod species: Acartia tonsa. The goal of this project was to assess the feasibility of establishing and maintaining cultures of Acartia tonsa, as well as to refine current culturing and egg separation methods. Initial culture methodology proved difficult for consistent production of eggs and collection of nauplii. The development of an airlift system for the separation of eggs from nauplii and adults, based on size, successfully increased the availability of eggs, nauplii and adults. The sensitivity and relative conditions of the copepod species was assessed by running a series of 48h acute toxicity tests with the reference toxicants 3,5-dichlorophenol, 4,4’-methylenebis(2,6-di-tert-butylphenol. The acute 48 hour median lethal dose concentration (LC50), the no observed effect concentration (NOEC), and the lowest observed effect concentration (LOEC) was analyzed for the three reference compounds for of A. tonsa.

Zooplankton are the major primary consumers in pelagic ecosystems, providing the principal pathway for energy transfer from primary production to higher trophic levels. The marine copepod Calanus finmarchicus is an important component of the pelagic food web in the North Atlantic and peripheral ecosystems, and forms an essential dietary component of a number of commercially important fish. As part of its life cycle, many C. finmarchicus overwinter in a diapause phase (a dormant overwintering phase where development is suppressed in adaptation to the seasonal food supply) at depths of 500 to 2000 m, but little is known about the triggers that initiate and terminate diapause, or the internal processes associated with these triggers. Understanding these processes is important, given that subtle changes in the environmental conditions which may affect diapause could have consequences for the entire Calanus-based ecosystem. In this study I took advantage of relatively easy access to a deep (> 100 m), isolated population of C. finmarchicus in Loch Etive (a sea loch on the west coast of Scotland) to sample Calanus finmarchicus monthly between April 2006 and June 2007 and measure lipid dynamics and gene expression associated with diapause. Chapter 1 of this thesis provides a general introduction to diapause and Calanus finmarchicus, Chapter 2 reports on the population of C. finmarchicus in Loch Etive, Chapter 3 reports changes in the lipid class and fatty acid composition of individual copepods, Chapter 4 reports on differential gene expression between diapausing and active C. finmarchicus and Chapter 5 provides a general discussion and puts this research into context. This study provides some initial insight into possible gene expression patterns, but further work is needed to attribute specific gene expression patterns with initiation and termination of diapause.

In the current study, behavioral responses of several species of calanoid copepods to mimics of oceanographic structure were observed and evaluated in the context of foraging and aggregation. Zooplankton distributions in oceanic habitats are often attributed to physical forcing; however, physical factors only act to drive ecological patterns at large scales (m to km). At fine to intermediate scales (cm to m) zooplankton behavior is believed to govern observed distributions, but the mechanisms and ecological significance of these behaviors are not well understood. In a water column, biological activity is often concentrated into one or a few regions, called thin layers, on the order of a meter thick, and zooplankton, such as copepods, must be able to reliably locate and exploit these patches for survival. Thin layers commonly are associated with oceanic structure such as flow gradients, fluid density jumps, or chemical composition gradients. Utilization of mechanosensory or chemosensory cues associated with thin layers may increase foraging success, thus translating into a significant ecological advantage. A laboratory apparatus was developed to create isolated and combined thin layer properties. Copepods then were exposed to laboratory mimics of thin layers. All of the tested species of copepods exhibited behavioral responses associated with area-restricted search behavior to one of the physical gradients (flow velocity or fluid density), but not both. Similar responses were observed for chemical exudate layer experiments and included increased proportional residence times, swimming speeds, and turn frequency. Food layers induced feeding responses from all tested species (increased proportional residence time, decreased swimming speed). Responses to various combinations of gradients were not fully synergistic, but suggested that some copepods employ a cue hierarchy to locate food-rich areas. Velocity or density gradients acted as initial cues for narrowing search regions, while chemical exudates induced responses that strengthened or removed the initial reactions. A simple foraging model was used to illustrate how such behavioral changes can lead to observed aggregations at larger temporal and spatial scales. Consequently, these results suggest that individual responses to oceanographic structure may have far reaching influence on population dynamics, succession, and biodiversity in coastal and pelagic ecosystems.

Locomotion is a key characteristic of almost all forms of life and is often accomplished, whether on land, in water, or in the air, by reciprocal motion of two or more appendages. Among the zooplankton, many species propel themselves by rhythmically beating multiple pairs of closely spaced leg-like appendages in a back-to-front (metachronal) pattern. The focus of this study is to understand the mechanical design, kinematic operation, and hydrodynamic result of metachrony in the zooplankton. In the first part of this study, Antarctic krill (Euphausia superba) are investigated as an ecologically important model species that metachronally beats its swimming legs (pleopods) to perform drag-based propulsion. Based on high speed videos of freely swimming Antarctic krill, hovering, fast forward swimming, and upside down swimming are identified as three distinct swimming modes with significantly different stroke amplitudes and beat frequencies. When transitioning between hovering and fast forward swimming, Antarctic krill first increase beat amplitude and secondarily increase beat frequency. In considering the design components that contribute to metachrony being a successful swimming technique, a comparison among many different species shows that the ratio between the appendage separation distance and appendage length is limited to a narrow range of values (i.e. 0.2 - 0.65). In the second part of this study, metachrony is examined at smaller length and time scales by examining the impulsive escape jump of a calanoid copepod (Calanus finmarchicus). The wake generated by the copepod's metachronally beating swimming legs is experimentally measured using a novel (and newly developed) tomographic particle image velocimetry (PIV) system capable of making volumetric 3D velocity measurements with high temporal and spatial resolution using IR illumination. The flow generated by the escaping copepod consisted of a stronger posterior vortex ring generated by the metachronally stroking swimming legs and a weaker one generated anteriorly around the body by the impulsive start of the escape, both of which decayed over time. The experiments also revealed azimuthal asymmetry in the vortices caused by body yawing and the action of the swimming legs, flow features not considered in previous axisymmetric computational and theoretical models of copepod jumps. While not accounting for this asymmetry, an impulsive stresslet is nonetheless useful in modeling the flow created by the escaping copepod and represents the flow more accurately than an impulsive Stokeslet. In the final part of this study, the flow associated with metachronal hovering in Antarctic krill is experimentally and theoretically investigated in regards to the energy requirements of the pelagic lifestyle. Volumetric flow measurements of a hovering Antarctic krill show that each stroking pleopod drags flow behind it such that a downward stream develops medially. The lateral exopodites induce tip vortices which add to the lift force on each appendage. Furthermore, the flow beneath the hovering krill develops into a pulsed jet with a Strouhal number in the 'high-efficiency zone' of 0.2 < St < 0.4. Actuator disk theory is used to make theoretical estimates of the induced power necessary to hover, the results of which match induced power values calculated from measured flow gradients contributing to viscous energy dissipation.

The focus of my thesis research is on the interplay between individual behavior, population dynamics and community-level processes within zooplankton communities in coastal Maine. The target organisms of my thesis work are marine copepods. Copepods are small (1-10 mm) crustaceans that perform the essential ecosystem function of consuming and assimilating primary production (phytoplankton) making it available to higher trophic levels such as commercially important fishes. Therefore, copepod population growth is of critical importance to marine food webs. Fertilization limitation has been suggested as a constraint on copepod population growth but field surveys describing the prevalence of fertilization limitation are lacking. During my doctoral research, I explored the in situ fertilization success of two marine copepod species, Temora longicornis and Eurytemora herdmani in coastal Maine. I collected monthly zooplankton samples and analyzed clutches from field-caught females using an egg-staining technique. My results indicate that both species exhibit fertilization limitation in nature and the factors correlated with their fertilization span population, community and ecosystem level factors. To determine a causal relationship between predator density and copepod mating success, I conducted laboratory experiments to assess the effects of a common mysid shrimp predator, Neomysis americana on Eurytemora herdmani mating success. I subjected males and females to predators or predator cues. I found that the presence of a mysid predator, or only a predator cue, reduced copulation frequency and spermatophore transfer leading to a 38-61% decrease in E. herdmani nauplii production. These results suggest that mysid predators can constrain copepod population growth through non-consumptive processes. To determine the effects that resources can impose on copepod behavior, I explored the behavioral and fitness consequences of Temora longicornis ingesting Alexandrium fundyense, a phytoplankton species that forms harmful algal blooms in coastal Maine. My results suggest that ingesting A. fundyense causes copepods to swim faster and with more directional persistence compared to control algae. Temora longicornis increased their average swimming velocity by 24%, which leads to a 24-54% increase in their theoretical encounter rate with predators. Therefore, these findings suggest behaviorally mediated copepod-algal interactions may have significant impacts on harmful algal bloom dynamics and the fate of toxins in marine food webs.

Although food quality is thought to play an important role in the survival of marine copepods, the extent of natural variability in food quality remains poorly characterized. Here I characterize the different scales at which food quality varies in copepods of the Strait of Georgia, British Columbia, Canada. Significant interannual variability occurs in the diet of Neocalanus plumchrus in the Strait of Georgia. Between 2001-06 the fatty acid profiles of N. plumchrus switched from omnivorous, oceanic signatures to herbivorous, diatom-dominated signatures. An index of food quality (DHA/EPA) is strongly correlated to the abundance of diapausing N. plumchrus, suggesting that the relative proportion of essential fatty acids provided by dinoflagellates and diatoms are related to the survival of this species. Combined fatty acid and stable isotope analysis indicated that the spring calanoid copepods of the Strait of Georgia occupy three trophic positions: Eucalanus bungii is herbivorous, Calanus marshallae and N. plumchrus are omnivorous, while Euchaeta elongata is carnivorous. Oceanic conspecifics of Strait of Georgia copepods experience a more omnivorous diet, as indicated by the presence of higher proportions of flagellate and carnivory markers, and lower proportions of diatom-based markers in their fatty acids. Despite spatial differences in the quality of their diets, the relative trophic positions of these copepods are constant as indicated by their stable isotope signatures. There is a correlation between the trophic information provided by stable isotopes and fatty acids. However, stable isotopes are not sensitive enough to capture the range of dietary variability observed in fatty acids, and fatty acids do not always provide reliable markers of carnivory and trophic position. Over the span of a season, copepods can utilize a wide range of dietary items including diatoms, flagellates, bacteria, detritus and microzooplankton. Copepods can switch from herbivory to carnivory in response to declining chlorophyll concentrations after the spring bloom, and are occasionally able to utilize detrital and bacterial sources. I conclude that the quality of copepod diets in the SoG varies on interannual, interspecific and seasonal scales. The implications of these findings are discussed in relation to ecosystem models of the area, and to copepod physiology.

Although food quality is thought to play an important role in the survival of marine copepods, the extent of natural variability in food quality remains poorly characterized. Here I characterize the different scales at which food quality varies in copepods of the Strait of Georgia, British Columbia, Canada. Significant interannual variability occurs in the diet of Neocalanus plumchrus in the Strait of Georgia. Between 2001-06 the fatty acid profiles of N. plumchrus switched from omnivorous, oceanic signatures to herbivorous, diatom-dominated signatures. An index of food quality (DHA/EPA) is strongly correlated to the abundance of diapausing N. plumchrus, suggesting that the relative proportion of essential fatty acids provided by dinoflagellates and diatoms are related to the survival of this species. Combined fatty acid and stable isotope analysis indicated that the spring calanoid copepods of the Strait of Georgia occupy three trophic positions: Eucalanus bungii is herbivorous, Calanus marshallae and N. plumchrus are omnivorous, while Euchaeta elongata is carnivorous. Oceanic conspecifics of Strait of Georgia copepods experience a more omnivorous diet, as indicated by the presence of higher proportions of flagellate and carnivory markers, and lower proportions of diatom-based markers in their fatty acids. Despite spatial differences in the quality of their diets, the relative trophic positions of these copepods are constant as indicated by their stable isotope signatures. There is a correlation between the trophic information provided by stable isotopes and fatty acids. However, stable isotopes are not sensitive enough to capture the range of dietary variability observed in fatty acids, and fatty acids do not always provide reliable markers of carnivory and trophic position. Over the span of a season, copepods can utilize a wide range of dietary items including diatoms, flagellates, bacteria, detritus and microzooplankton. Copepods can switch from herbivory to carnivory in response to declining chlorophyll concentrations after the spring bloom, and are occasionally able to utilize detrital and bacterial sources. I conclude that the quality of copepod diets in the SoG varies on interannual, interspecific and seasonal scales. The implications of these findings are discussed in relation to ecosystem models of the area, and to copepod physiology.

Made available in DSpace on 2015-04-17T14:55:25Z (GMT). No. of bitstreams: 1 arquivototal.pdf: 1509338 bytes, checksum: e0a2529155ec99b45b55b847965af181 (MD5) Previous issue date: 2012-07-25 / Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - CAPES / Cyclopoid copepods have been used as biological agents in successful programs to control mosquito larvae, but the impacts of this predation on adult mosquito populations are still poorly understood. The present study compared the sex ratios and body sizes (measure as wing length) of Aedes albopictus mosquitoes emerging from recipients containing the copepod predator Mesocyclops ogunnus with control situations without this predator. We found that copepod predation significantly biased mosquito sex ratios towards females, and that both the males and females emerging from copepod-containing recipients were significantly larger than control insects. The ecological and epidemiological consequences of the changes induced by copepod predation on mosquito populations are discussed. / Copépodos ciclopóides têm sido utilizados como agentes de controle biológicos em programas de controle de larvas de mosquito, mas os impactos dessa predação sobre as populações de mosquitos adultos ainda são pouco compreendidos. O presente estudo comparou as razões sexuais e o tamanho do corpo de mosquitos Aedes albopictus (com base no comprimento da asa) que emergiram de recipientes contendo o copépodo predador Mesocyclops ogunnus com aqueles que emergiram de recipientes controles. Com base nos resultados obtidos constatou-se que a predação de copépodos alterou significativamente a razão sexual de mosquitos favorecendo fêmeas. Machos e fêmeas que emergiram de recipientes contendo copépodo foram significativamente maiores em comparação aqueles que emergiram do controle. As alterações induzidas pela predação de copépodos em populações de mosquitos apresentam importâncias ecológicas e epidemiológicas, visto serem as fêmeas de mosquitos do gênero Aedes os principais vetores do dengue .