Metacommunity Dynamics Of Benthic Macroinvertebrates In A Large River

Metacommunity ecology focuses on the interaction between local communities and is inherently linked to dispersal as a result. Within this framework, communities are structured by a combination of in-site responses to the immediate environment (species sorting), stochasticity (patch dynamics), and connections to other communities via distance between communities and dispersal (neutrality), and source-sink dynamics (mass effects; see Chapter 1 for a detailed description of metacommunity theory, the study site, and macroinvertebrate communities found). In Chapter 2 I describe spatial scale of study and dispersal ability as both have the ability to influence the degree to which communities interact. However, little is known about how these factors influence the importance of all metacommunity dynamics. I compared dispersal mode of immature aquatic insects and dispersal ability of winged adults across multiple spatial scales in a large river. The strongest drivers of river communities were patch dynamics, followed by species sorting, then neutrality. Active dispersers during aquatic lifestages on average exhibited lower patch dynamics, higher species sorting, and significant mass effects compared to passive dispersers. Active and strong dispersers also had a scale-independent influence of neutrality, while neutrality was stronger at broader spatial scale for passive and weak dispersers. These results indicate as dispersal ability increases patch dynamics decreases, species sorting increases, and neutrality should decrease. The perceived influence of neutrality may also be dependent on spatial scale and dispersal ability. In Chapter 3 I describe how river benthic macroinvertebrate communities may influence tributary invertebrate communities via adult flight and tributaries may influence mainstem communities via immature drift. This relationship may also depend on relative mainstem and tributary size, as well as abiotic tributary influence on mainstem habitat. To investigate the interaction between a larger river and tributary I sampled mainstem benthic invertebrate communities and quantified habitat of a 7th order river (West Branch Susquehanna River) above and below a 5th order tributary confluence, as well as 0.95-3.2 km upstream in the tributary. Non-metric multidimensional scaling showed similar patterns of clustering between sampling locations for both habitat characteristics and invertebrate communities. In addition, mainstem river communities and habitat directly downstream of the tributary confluence cluster tightly together, intermediate between tributary and mid-channel river samples. In Bray-Curtis dissimilarity comparisons between tributary and mainstem river communities the furthest upstream tributary communities were least similar to river communities. Middle tributary samples were also closest by Euclidean distance to the upstream mainstem riffle and exhibited higher similarity to mid-channel samples than the furthest downstream tributary communities. My results indicate river and tributary benthic invertebrate communities may interact and likely result in direct and indirect mass effects of a tributary on the downstream mainstem community by invertebrate drift and habitat restructuring via material delivery from the tributary. I also showed likely direct effects of adult dispersal from the river and oviposition in proximal tributary locations where Euclidian, rather than river, distance may be more important in determining river-tributary interactions.

Tributary and Mainstem Benthic Macroinvertebrate Communities Linked By Direct Dispersal and Indirect Habitat Alteration

Benthic communities in tributary-mainstem networks might interact via downstream drift of invertebrates or material from tributaries and adult dispersal from the mainstem. Depending on the strength of these interactions, mainstem downstream communities are expected to be more similar to tributary communities due to drift or habitat alteration. Communities not connected by flow are expected to be similar due to adult dispersal but decreasing in similarity with distance from the mainstem. We investigated interactions between invertebrate communities of a 7th order river and 5th order tributary by comparing benthic community structure in the river upstream and downstream of the tributary confluence and upstream in the tributary. Non-metric multidimensional scaling showed invertebrate communities and habitat traits from river locations directly downstream of the tributary clustered tightly, intermediate between tributary and mid-channel river locations. In addition, Bray-Curtis dissimilarity increased between the mainstem and tributary with distance upstream in the tributary. Our results indicate that similarities between mainstem and tributary communities are potentially caused by direct mass effects from tributary to downstream mainstem communities by invertebrate drift and indirect mass effects by habitat restructuring via material delivery from the tributary, as well as potential effects of adult dispersal from the river on proximal tributary communities.

Ecology of larval fishes and large zooplankton in the Keweenaw Current region of Lake Superior, with special focus on lake herring, Coregonus artedi

I assessed the influence of the Keweenaw Current and spring thermal bar on the distribution of larval fishes and large zooplankton in Lake Superior. In 1998 and 1999, samples were collected from inshore (0.2 – 3.0 km from shore) and offshore (5.0 – 9.0 km from shore) locations on three transects off the western coast of the Keweenaw Peninsula, Michigan. For larval fishes, density and size distribution patterns of lake herring (Coregonus artedi), rainbow smelt (Osmerus mordax), burbot (Lota lota), deepwater sculpin (Myoxocephalus thompsoni), and spoonhead sculpin (Cottus ricei) suggest a seasonal inshore to offshore movement. For zooplankton, seasonal warming appeared to be the major factor that limited planktonic catches of the primarily benthic Mysisrelicta and Diporeia spp., while simultaneously stimulated growth and reproduction of the cladocerans Daphnia spp., Holopedium gibberum, and Bythotrephes cederstroemi. In contrast, calanoid copepods as a group were abundant throughout the entire sampling season. The greatest abundances of zooplankton were generally encountered offshore, even for the cladocerans, which apparently expanded from inshore to offshore locations with seasonal warming. In 2000, sampling efforts focused on lake herring. Samples were collected from surface waters at 0.1 – 17.0 km from shore on two transects. Lake herring larvae were also reared in the laboratory from eggs in order to validate the use of otolith microstructure for aging. Increment deposition was not statistically different from a daily rate starting from 28 days after hatching, near the time of yolk-sac absorption, but larvae with lower growth rates could not be aged as accurately. In Lake Superior, lake herring tended to be slightly more abundant, larger, and older at inshore locations, but a dense patch of younger larvae was also encountered 7 – 13 km from shore. The distribution iiipatterns suggest that larvae were transported by prevailing currents into the study region, possibly from the more productive spawning regions in western Lake Superior. Growth rates were suppressed at offshore locations where temperatures were less than 8°C. These results indicate that lake herring larvae may be transported far distances from spawning concentrations by longshore currents, and water temperatures may largely control their growth.

Abundance and biomass of the benthic infauna of the North Sea

In 1986 participants of the Benthos Ecology Working Group of ICES conducted a synoptic mapping of the infauna of the southern and central North Sea. Together with a mapping of the infauna of the northern North Sea by Eleftheriou and Basford (1989, doi:10.1017/S0025315400049158) this provides the database for the description of the benthic infauna of the whole North Sea in this paper. Division of the infauna into assemblages by TWINSPAN analysis separated northern assemblages from southern assemblages along the 70 m depth contour. Assemblages were further separated by the 30, 50 m and 100 m depth contour as well as by the sediment type. In addition to widely distributed species, cold water species do not occur further south than the northern edge of the Dogger Bank, which corresponds to the 50 m depth contour. Warm water species were not found north of the 100 m depth contour. Some species occur on all types of sediment but most are restricted to a special sediment and therefore these species are limited in their distribution. The factors structuring species distributions and assemblages seem to be temperature, the influence of different water masses, e.g. Atlantic water, the type of sediment and the food supply to the benthos.

Bioturbation determines the response of benthic ammonia-oxidizing microorganisms to ocean acidification

Ocean acidification (OA), caused by the dissolution of increasing concentrations of atmospheric carbon dioxide (CO2) in seawater, is projected to cause significant changes to marine ecology and biogeochemistry. Potential impacts on the microbially driven cycling of nitrogen are of particular concern. Specifically, under seawater pH levels approximating future OA scenarios, rates of ammonia oxidation (the rate-limiting first step of the nitrification pathway) have been shown to dramatically decrease in seawater, but not in underlying sediments. However, no prior study has considered the interactive effects of microbial ammonia oxidation and macrofaunal bioturbation activity, which can enhance nitrogen transformation rates. Using experimental mesocosms, we investigated the responses to OA of ammonia oxidizing microorganisms inhabiting surface sediments and sediments within burrow walls of the mud shrimp Upogebia deltaura. Seawater was acidified to one of four target pH values (pHT 7.90, 7.70, 7.35 and 6.80) in comparison with a control (pHT 8.10). At pHT 8.10, ammonia oxidation rates in burrow wall sediments were, on average, fivefold greater than in surface sediments. However, at all acidified pH values (pH < = 7.90), ammonia oxidation rates in burrow sediments were significantly inhibited (by 79-97%; p < 0.01), whereas rates in surface sediments were unaffected. Both bacterial and archaeal abundances increased significantly as pHT declined; by contrast, relative abundances of bacterial and archaeal ammonia oxidation (amoA) genes did not vary. This research suggests that OA could cause substantial reductions in total benthic ammonia oxidation rates in coastal bioturbated sediments, leading to corresponding changes in coupled nitrogen cycling between the benthic and pelagic realms.

pH and calcium change in the microenvironment of a benthic foraminifer (Ammonia sp.) and its size during experiments

Calcareous foraminifera are well known for their CaCO3 shells. Yet, CaCO3 precipitation acidifies the calcifying fluid. Calcification without pH regulation would therefore rapidly create a negative feedback for CaCO3 precipitation. In unicellular organisms, like foraminifera, an effective mechanism to counteract this acidification could be the externalization of H+ from the site of calcification. In this study we show that a benthic symbiont-free foraminifer Ammonia sp. actively decreases pH within its extracellular microenvironment only while precipitating calcite. During chamber formation events the strongest pH decreases occurred in the vicinity of a newly forming chamber (range of gradient about 100 µm) with a recorded minimum of 6.31 (< 10 µm from the shell) and a maximum duration of 7 h. The acidification was actively regulated by the foraminifera and correlated with shell diameters, indicating that the amount of protons removed during calcification is directly related to the volume of calcite precipitated. The here presented findings imply that H+ expulsion as a result of calcification may be a wider strategy for maintaining pH homeostasis in unicellular calcifying organisms.