The aim of the present study was to assess the ecological role of the euthecosome pteropod, Limacina retroversa, in particular, and the mesozooplankton community, in general, in the pelagic ecosystem of the Polar Frontal Zone (PFZ), Southern Ocean. Data were collected from four oceanographic surveys to the Indian sector of the PFZ during austral autumn 2000, 2002, 2004 and 2005. Copepods, mainly Calanus simillimus, Oithona similis, Clausocalanus spp. and Ctenocalanus spp., typically dominated total mesozooplankton counts, accounting for, on average, between 75.5 % and 88.1 % (Mean = 77.4 %; SD = 13.4 %) of the total, during the present investigation. Results of the study indicate that L. retroversa may, at times, contribute substantially to total mesozooplankton abundances. During the study, L. retroversa contributed between 0.0 and 30.0 % (Mean = 5.3 %; SD = 7.1 %) to total mesozooplankton numbers. Significant small-scale variability in abundance and size structure of L. retroversa and abundance of copepods was minimal. Inter-annual variability, on the other hand, was significant between some years. Total pteropod numbers were greatest during April 2002 and 2004, while copepods exhibited greatest abundances during April 2004 only. Pearson’s Correlation analysis suggested that L. retroversa abundances were positively correlated to total surface chlorophyll-a (chl-a) concentrations. The significantly lower chl-a concentrations recorded during April 2005 may explain the reduced pteropod numbers observed during that survey. The size class structure of L. retroversa comprised mainly small and mediumsized individuals during all four surveys. This corresponds well with records from the northern hemisphere (sub-Arctic and Arctic waters) where Limacina spp. are reported to exhibit maximum spawning during mid to late-summer. Higher abundances of large individuals only occurred during April 2005, when chl-a concentrations were very low; possibly the result of delayed spawning, due to reduced food availability. Ingestion rates of the four most abundant copepods, determined using the gut fluorescence technique, ranged between 159.32 ng (pigm) ind⁻¹ day⁻¹ and 728.36 ng (pigm) ind⁻¹ day⁻¹ (Mean = 321.01 ng (pigm) ind⁻¹ day⁻¹; SD = 173.91 ng (pigm) ind⁻¹ day). Ingestion rates of L. retroversa were much higher, ranging from an average of 4 28.68 ng (pigm) ind⁻¹ day⁻¹ in April 2002 to 4 196.88 ng (pigm) ind⁻¹day⁻¹in April 2005 (Mean = 4157.36 ng (pigm) ind⁻¹ day⁻¹; SD = 35.37 ng (pigm) ind⁻¹day⁻¹). Average daily grazing rates for the pteropod varied between 0.39 mg (pigm) m⁻² day⁻¹ in April 2005 and 17.69 mg (pigm) m-2 day-1 in April 2004 (Mean = 6.13 mg (pigm) m⁻² day⁻¹; SD = 11.04 mg (pigm) m⁻² day⁻¹); corresponding average daily grazing impacts ranged between 8.4 % and 139.8 % of the phytoplankton standing stock in April 2005 and 2004, respectively (Mean = 48.5 %; SD = 84.5 %). Average daily grazing rates of the four copepods ranged from 4.58 mg (pigm) m⁻² day⁻¹ to 8.77 mg (pigm) m⁻² day⁻¹ -1, during April 2002 and 2004, respectively (Mean = 6.28 mg (pigm) m⁻² day⁻¹; SD = 5.94 mg (pigm) m⁻² day⁻¹). Collectively, the copepods removed an average of between 31.6 % and 89.8 % of the phytoplankton standing stock per day, during April 2002 and 2004, respectively (Mean = 70.8 %; SD = 86.7 %). The daily grazing impact of the copepods accounted for an average of between 40.4 % and 87.8 % of the total zooplankton grazing impact, during April 2004 and 2005, respectively (Mean = 75.0 %; SD = 65.5 %). L. retroversa was responsible for an average of 52.4 % and 59.5 % of the total zooplankton grazing impact, during April 2002 and 2004, respectively. However, during April 2005, when L. retroversa numbers were significantly lower than previous years, the pteropod contributed an average of only 7.5 % to the total zooplankton grazing impact. Thus, during the present investigation,the pteropod was responsible for removing a mean of 48.9 % of the available phytoplankton (SD = 74.9 %). The predation impact of the dominant carnivorous macrozooplankton and micronekton in the PFZ was determined during April 2004 and 2005 using daily ration estimates obtained from the literature. Additionally, gut content analysis was used to determine the contribution of L. retroversa to the diet of the dominant predators. Average predation impact ranged from 1.1 % and 5.7 % of the total mesozooplankton standing stock during April 2004 and 2005, respectively (Mean = 3.8 %; SD = 12.3 %). Chaetognaths and euphausiids dominated total carnivore numbers and made the greatest contributions to total predation impact during both years. Copepods appeared to be the main prey item of the dominant carnivorous macrozooplankton-micronekton in the region. L. retroversa was only detected in the gut contents of the amphipod, Themisto gaudichaudi, but not in either of the chaetognath species (Eukrohnia hamata and Sagitta gazellae) or the myctophid fish (Electrona spp.). The pteropod was found in 19 % of amphipod guts dissected. Pearson’s Correlation analyses showed that the four major predatory zooplankton groups found in the PFZ (chaetognaths, euphausiids, amphipods and myctophid fish) were positively correlated to abundances of L. retroversa, suggesting that the pteropod might be an important prey item for many of the carnivorous macrozooplankton/micronekton in the PFZ. To conclude, L. retroversa may play an important role in the pelagic ecosystem of the PFZ, in austral autumn. However, ocean acidification and calcium carbonate undersaturation (as a result of increased anthropogenic carbon dioxide emissions), that is predicted to occur within the next 50 – 100 years, will most likely have significant implications for the Sub-Antarctic pelagic ecosystem if L. retroversa cannot adapt quickly enough to the changes.

The trophodynamics of the numerically dominant mesozooplankton (200-2000 m) in the vicinity of the Subtropical Convergence (STC) in the Indian sector of the Southern Ocean during austral autumn (April / May) 2007 were investigated as part of the Southern Ocean Ecosystem Variability Study. The survey consisted of six north-south transects each bisecting the STC between 38º to 43ºS and 38º to 41º45’E. In total, 48 stations situated at 30 nautical mile intervals were occupied over a period of ten days. Hydrographic data revealed a well defined surface and sub-surface expression of the STC, which appeared to meander considerably between 41ºS and 41º15’S. Surface chlorophyll-a (chla) concentrations were low, ranging between 0.08 and 0.68 mg chl-a.m-3 and were generally dominated by the picophytoplankton (<2 m) which made up 66.6% (SD±17.6) of the total pigment. Chl-a concentrations integrated over the top 150m of the water column ranged between 11.97 and 40.07 mg chl-a.m-2 and showed no significant spatial patterns (p>0.05). Total integrated mesozooplankton abundance and biomass during the study ranged between 3934.9 and 308521.4 ind.m-2 (mean = 47198.19; SD±62411.4 ind.m-2) and between 239.8 and 4614.3 mg Dwt.m-2 (mean = 1338.58; SD ±1060.5), respectively. Again, there were no significant spatial patterns in the total mesozooplankton abundance or biomass within the region of study (p>0.05). No significant correlations were found between biological (chlorophyll-a concentrations and zooplankton abundance) and physico-chemical variables (temperature and salinity) (p>0.05). The total mesozooplankton community was numerically dominated by copepods of the genera Pleuromamma, Calanus, Oncaea and Oithona. Other important representatives of the mesozooplankton community included the tunicate, Salpa thompsoni, and the pteropod, Limacina retroversa. At the 40% similarity level, numerical analysis identified five distinct mesozooplankton groupings within the survey area. Differences between the groupings were associated with changes in the relative contribution of numerically dominant species rather than the presence or absence of individual species. No groupings were associated with any specific feature of the front within the survey area. The feeding rates of the six most numerically abundant mesozooplankton species (Calanus simillimus, Limacina retroversa, Pleuromamma abdominalis, Clausocalanus breviceps, Oncaea conifera, Salpa thompsoni) accounting for on average 39% of the total mesozooplankton counts, were investigated using the gut fluorescence technique. For all species, the total gut pigment contents during the night time were significantly higher than the daytime values (p<0.05 for all species). The gut evacuation rates (k) for selected mesozooplankton ranged between 0.14 and 0.81 h-1. The ingestion rates ranged between 147.8 and 5495.4 ng(pigm)ind-1.day-1 which corresponded to a daily ration of between 2.4 and 10.9% body carbon. The combined grazing impact of the selected species on the daily phytoplankton standing stock was highly variable and ranged between 1.2 and 174.1% with an average of 27.3% (SD±38.78%) within the survey area. The highest grazing impact (>60%) was typically associated with those stations where the pteropod, L. retroversa, and the tunicate, S. thompsoni, contributed more than 5% of the total mesozooplankton counts. No significant differences were found in the grazing impact of any or all selected species situated either north, south or in the immediate vicinity of the front (p>0.05 in all cases). The lack of defined spatial patterns in the mesozooplankton abundance and community structure suggests that the STC did not act as a significant biogeographic barrier to the distribution of mesozooplankton during the study. It is presumed that the large scale mixing event caused by a storm prior to this study was responsible for the observed lack of elevated biological activity within the region of the STC.

Despite high densities in an upstream lake, the herbivorous calanoid copepod, Diaptomus leptopus, is extremely rare in oligotrophia Eunice Lake in the Coastal Range Mountains near Vancouver, British Columbia. In situ experiments conducted in 1979 and 1980 tested the hypothesis that competition from zooplankton species resident in Eunice Lake prevents immigrant D. leptopus from colonizing the lake. Polyethylene enclosures, each holding 29,000 1 of lake water, contained all experimental treatments. Experiments in 1979 exposed a standard density of D. leptopus to all Eunice Lake zooplankton species at lake densities (control), to all Eunice Lake species at reduced densities (low density), and to all Eunice Lake species except one of Daphnia rosea (Daphnia-removal), Diaptomus kenai (kenai-removal), or Diaptomus tyrelli (tyrelli-removal). Improved performances of D. leptopus in non-control treatments was evidence for competition in controls. Measures of performance included density, number of eggs per female, and adult size. Similarity between D. leptopus performances in control, Daphnia-removal, and kenai-removal treatments plus relatively high concentrations of D. leptopus nauplii in the tyrelli-removal treatment suggested that D. leptopus did not compete with species other than D. tyrelli. However, similarity between tyrelli-removal and low density treatments in concentrations of D. leptopus nauplii confounded competition between D. leptopus and D. tyrelli with diffuse competition from several Eunice Lake species combined. D. leptopus overlapped in vertical distribution and seasonal cycle more with D. tyrelli than with other species. Differences' among experimental treatments in algal size compositions did not conclusively show whether zooplankton partitioned food resources. Experiments in 1980, designed to separate D. tyrelli competition from diffuse competition, did not provide any evidence of interspecific interactions. By increasing zooplankton metabolic efficiencies, cool temperatures in 1980 may have virtually eliminated, competition for algal foods. / Science, Faculty of / Zoology, Department of / Graduate

Magister Scientiae (Biodiversity and Conservation Biology) / 2022-01-31

This thesis represents a zooplankton study of Tomales Bay, Marin County, California. It consists of two papers. The first paper, "The Abundance of Copepods in Tomales Bay, California" has been submitted to the California Fish and Game for publication. The second paper, "Diet Studies of the Ctenophore Pleurobrachia bachei has been submitted to Fishery Bulletin. The raw data has been compiled in the Appendix.

In lakes, fish and zooplankton can be both sources and sinks of nitrogen (N) and phosphorus (P) through the consumption of organic N and P, and subsequent excretion of bioavailable inorganic forms. These source/sink dynamics, known as consumer-driven nutrient recycling (CNR), may, in turn, control the availability of potentially limiting nutrients for algal primary production. In this dissertation I investigate the importance and controls of CNR as a source of inorganic N and P for primary production (Chapter 2). I then examine zooplankton CNR as a mechanism for increasing nutrient mean resident time (MRT) in the mixed layer of lakes (Chapter 3). Finally, I assess whether zooplankton communities dominated by different taxa can affect N versus P deficient conditions for phytoplankton production through differential N and P recycling rates (Chapter 4). Direct excretion of N and P by fish communities was modest in arctic lakes, and accounted for < 4 % of the N and P required for primary production. Recycling of N and P by zooplankton communities was relatively high, and the fraction of algal N and P demand supplied by zooplankton CNR ranged from 4 - 90% for N and 7 - 107% for P. MRT of 15N, measured in the mixed layer of an arctic lake, was ~16 days, compared to 14 days predicted by a ecosystem model simulation with zooplankton N recycling and 8 days in a model simulation where zooplankton N recycling was absent. The 75% increase in N MRT between model simulations with and without zooplankton recycling suggests that zooplankton N recycling is an important mechanism for retaining N in lake ecosystems. I observed relatively high negative correlations between precipitation and phytoplankton N (r = -0.33) and P (r = -0.30) deficiencies. I also observed a significant positive correlation (r = 0.42, p = 0.03) between zooplankton communities with higher copepod biomass, relative to cladoceran biomass, and phytoplankton N-deficient conditions. These results suggest that when precipitation is high N and P deficiency is low in the phytoplankton. When precipitation is low, however, zooplankton communities composed primarily of copepods contribute to N-deficient conditions for phytoplankton production.

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This study examined the structural and functional development of zooplankton communities in 12 new experimental ponds for one year and tested four predictions derived from the Random Placement Hypothesis (Coleman 1981). Physico-chemistry, zooplankton colonization dynamics, zooplankton community structure and function were analyzed every two weeks from 5 February 1988 to 10 February 1989. Ponds varied in physico-chemistry at points in time but followed similar patterns during the study year. Ponds were not colonized by zooplankton similarly. Some species occurred in all ponds with about the same timing, but many species exhibited variable timing among ponds or never occurred in more than a few ponds. Colonization curves varied among ponds and through time, and species accrual curves differed in both accrual rates and the numbers of species accrued. Observed colonization curves did not closely match the curve expected according to the Random Placement Hypothesis. Zooplankton community structure also varied among ponds. Multivariate analyses could not discern similar trends in zooplankton community structure among ponds due to the disparity of species trends among ponds. species data were pooled into taxa (Copepoda, Cladocera, Rotifera, ostracoda and Chaoborus) and analyzed. Rotifers dominated zooplankton communities in densities and biomass, and ponds differed in taxa densities and biomasses. Zooplankton community function was more similar among ponds than community structure. Multivariate analyses indicated ponds generally followed similar trajectories in zooplankton community function through the year. Zooplankton did not colonize experimental ponds equally and did not develop similar zooplankton community structure among ponds. Dispersal processes probably limited colonization and development of zooplankton community structure. Zooplankton community function was generally more similar among ponds than community structure, probably due to the functional redundancy of zooplankton species. Implications of these results for experimental pond studies are discussed. / Ph. D.