Invertebrates influence primary productivity and nutrient cycling in ecosystems. They are also important links between producer and higher trophic levels. Despite their important role in terrestrial ecosystem function, invertebrates are frequently overlooked in ecological restorations. Thus, the objective of this research was to quantify how belowground macroinvertebrate ecosystem engineers and communities change over time following ecological restoration and examine whether the source of dominant plant species and the composition of non-dominant plant species influence aboveground macroinvertebrates community structure in restored prairie. A chronosequence design (space for time substitution) was used to determine the role of restoration age, plant community, and soil structure on the recovery of two belowground macroinvertebrate ecosystem engineers (Chapter 2). Ants and earthworms were sampled from cultivated fields, grasslands restored for 1-21 years, and native prairie. Earthworm abundance increased with time since cessation of cultivation, concomitant with prairie establishment. The abundance and biomass of ants were more related to the structure of restored plant communities than time since restoration. The dominance of exotic earthworms, and a generalist ant species in these restorations, coupled with their known capacity to alter soil properties and processes, may represent novel conditions for grassland development. The same chronosequence of agricultural fields, restored prairies, and prairies that were never cultivated was used to address the second objective of this research, which was to quantify how the belowground macroinvertebrate community composition changed in response to ecological restoration and whether the communities became representative of undisturbed (“target”) communities. Macroinvertebrate communities in the two remnant prairie sampled were distinct from restorations and continuouslycultivated fields. The macroinvertebrate communities in prairie that had never been cultivated were also distince from each other, indicating a “target” community is hard to define. Belowground macroinvertebrates changed in a trajectory that was not representative of either remnant prairie, but was representative of the an average of both remnant prairies. Thus, if you reconstruct prairie from cultivated soil conditions (“build it”), macroinvertebrates will colonize (“come”), but attaining a community representative of a specific target may require introduction from that target. Colonization of macroinvertebrates in restorations aboveground are most likely to be influenced by aspects of the plant community. A third objective of this research was to quantify whether variation in dominant species (cultivars vs. local ecotypes) and composition of subordinate species (local species pools) influence the composition of aboveground macroinvertebrates. Macronvertebrate abundance, richness, diversity, trophic groups and community composition in late summer did not vary between prairie restored with cultivar and local ecotypes of the dominant grasses. This was observed in two field experiments. The species pool treatment did influence the macroinvertebrate community, as one species pool had slightly higher morphospecies diversity and hymenopterans that the other two species pools. This was likely due to the presence of an ant-tended legume, Chamaecrista facsciculata Michx., in one species pool. Overall, this research demonstrates that time since the cessation of disturbance (cultivatation) and plant communities influence macroinvertebrate communities in restored prairie. Restored prairies in the Midwest are likely to be colonized by exotic earthworms and cosmopolitan ants. More research is needed to reveal how they influence ecosystem functioning.Belowground, macroinvertebrate communities may not represent restoration “targets” and these “targets” may be hard to define if remnants are rare or there is a high degree of spatial variation on the landscape. Variation in plant communities above ground appears to influence the structure of aboveground macroinvertebrate communities more than variation within dominant species. Whether this aboveground variation is reflected is reflected belowground deserves further investigation.

Low species diversity in the Arctic promotes strong food-web linkages, as changes in abundance of one species may influence many others. Using harvest records, I determined Arctic fox populations are declining in their southern distributional range due to shallower snow potentially limiting density of lemmings, their primary prey, which live and breed beneath snow. Additionally, warm fall and spring temperatures are shortening access to alternative prey, seals on sea ice. Arctic foxes also influence other species through non-trophic interactions, as lemming winter nests were found on 70% of fox dens examined. I determined warmer subnivean temperatures promoted by accumulation of thick snow leeward of tall vegetation on dens attracted lemmings to these dens. Furthermore, lemming reproduction was higher dens compared to traditional lemming habitat. This research highlights the impact of climatic variables on Arctic predator-prey interactions and the importance of understanding impacts of trophic and non-trophic interactions on species demographics. / February 2017

I estimated the non-trophic effects of arctic fox (Vulpes lagopus) denning activities on soil nutrient dynamics, vegetation production and quality, snow cover thickness, and their primary terrestrial prey, collared lemmings (Dicrostonyx richardsoni), near Churchill, Manitoba in April, June and August 2014. Arctic foxes increased soil inorganic nitrogen and extractable phosphorous concentration on their dens. This increase in soil nutrient levels resulted in greater vegetation quantity (measured as biomass and cover) and quality (measured as nitrogen content) on dens. Increased vegetation cover, specifically Salix sp. and Leymus mollis cover, positively affected snow cover thickness on dens by trapping blowing snow. Increased snow cover thickness made dens attractive nesting sites to collared lemmings (measured as lemming nest counts). In addition, dens with lemming nests had greater snow cover thickness compared to dens without lemming nests. Greater vegetation quantity and quality on dens could also attract lemmings to dens for winter nesting. / February 2016

Recent theoretical and technological advances in ecosystem science have dramatically expanded the ways in which scientists can pursue and explore ecological questions. For my thesis research, I integrated the recent theoretical concept of organisms as ecosystem engineers with the relatively recent development of stable isotope tracer tests to ask the question: how does the invasive common carp affect stream ecosystem structure and function? To investigate the structuring role of carp, I measured autotroph seasonal distribution and abundance and macroinvertebrate seasonal abundance and diversity within two stream reaches in Spring Creek, Utah, USA; one with low carp biomass (LCB) and one with high carp biomass (HCB). I installed a series of carp exclosures in the HCB reach to examine the response of the stream to carp exclusion. To explore the effects of carp on stream nitrogen dynamics, I performed a three week, continuous injection of 15N as ammonium chloride. The macrophyte and macroinvertebrate community was severely depauperate in the HCB reach compared to the LCB reach. The observed rapid colonization of a relatively abundant and diverse macrophyte and macroinvertebrate community at the carp exclusion sites in the HCB reach not only indicates that carp engineering reduces the abundance and diversity of these communities, but also highlights the importance of the spatial distribution of engineered and non-engineered patches in dictating the temporal scale of re-colonization. Carp engineering had a simplifying effect on stream N dynamics that ultimately limited the uptake and retention capacity of the HCB reach. For example, macrophytes played a dominant role in the N dynamics of the LCB reach by directly assimilating NH4, retaining N rich FBOM, and by providing habitat necessary to support an abundant and relatively diverse macroinvertebrate community that facilitated greater trophic transfer of nitrogen. Conversely, carp reduction of macrophytes in the HCB reach resulted in an overall reduction in areal uptake rates of NH4, reduced trophic transfer of N, and significantly reduced N retention. These results clearly indicate that carp engineering reduces macrophyte and macroinvertebrate abundance and diversity in streams and that N dynamics are simplified in carp engineered patches.

Marine ecosystems are increasingly being subject to human impact from diverse recreational and commercial activities, not necessarily restricted to those of a marine nature. This has significant implications for biodiversity. The large dog cockle, Tucetona laticostata, once occurred live in Rangitoto Channel, Hauraki Gulf, although this species no longer appears to occur there, most likely as a consequence of repeated dredging and channel excavation and continued siltation. Tucetona laticostata still occurs in a few isolated pockets of sea bed throughout Hauraki Gulf, particularly off Otata Island, part of the Noises complex of islands, where it resides partially buried in shell and rock gravels in shallow water (to 15 metres depth). The shells of T. laticostata collect in large post-mortem deposits in an area ramping from the sea bed off southwestern Otata Island. These mounds differ significantly in structural complexity from those of adjacent, extensively fragmented shell and rock gravels. Using the mounds of T. laticostata shell as a proxy for structural complexity, in order to appraise the effect of complexity on benthic-invertebrate diversity, the sea bed off southwestern Otata Island was sampled quarterly at two depths and in both T. laticostata shell mounds and adjacent extensively fragmented shell and rock gravels. These data were complemented with those from additional surveys around Otata Island, and off eastern Motutapu Island to determine the distribution and composition of benthic-invertebrate community assemblages throughout the region, and from concurrent surveys throughout the Waitemata Harbour and inner Hauraki Gulf to determine the current distribution of T. laticostata in this region. The number of benthic invertebrate species and individuals within T. laticostata habitat almost always was higher than that occurring within extensively fragmented shell- and rock gravel habitat, with densities to 142,385 individuals m-2 encountered. Temporal and spatial variations in benthic community structure also are reported for the two habitats, T. laticostata-based shells and extensively fragmented shell- and rock gravels. The numbers of species were higher amongst samples collected off the southwestern and eastern sides of Otata Island than elsewhere around this island, or of eastern Motutapu Island. Of the 351 species reported from all Otata and Motutapu Island samples combined, 73% of them occurred off southwestern Otata Island, 30% of which were found exclusively within T. laticostata shell habitat, and 10.5% within extensively fragmented shell and rock gravel habitat. The sea bed off southwestern Otata Island is regularly, seasonally dredged by recreational scallop fishers, in addition to being a popular small-vessel anchorage site. Both of these activities, dredging and anchorage, stand to reduce substratum complexity by fragmentation and dispersal of the valves of T. laticostata. Given the unique benthic invertebrates reported from T. laticostata shell deposits reported from southwestern Otata Island, any activity that damages the shells of this species, regardless of whether they are live or dead, is likely to result in loss of biodiversity. Admittedly, many of species identified as major contributors to differences in benthic invertebrate assemblages between T. laticostata shell-based habitats and those of extensively fragmented shell and rock gravels are not particularly charismatic or large, but each likely plays a role in local food webs and/or sediment and water column chemistry. It was not the intention of this research to determine the effects of anthropogenic disturbances like dredging or vessel anchorage on benthic-invertebrate communities off southwestern Otata Island. However, given the reported differences in species diversity within the structurally complex substratum provided by T. laticostata, conservation of biogenic reef-forming species like it might be a prudent, precautionary measure to take.

In arid landscapes limited resources are concentrated into fertile patches that allow greater productivity than would otherwise be expected. Fertile patches are created and maintained by processes that modify soil topography and interrupt the flow of resources across the landscape. These processes may be abiotic or biotic in origin. Species that modify, maintain or create habitat have been termed ecosystem engineers. One group of soil disturbing ecosystem engineers creates fertile patches by modifying soil microtopography through foraging for food and creating habitat. This thesis examines the effects of soil foraging animals on resource concentration in arid environments and how effects are moderated by species and landscape. Chapter 1 provides an overview of the importance of fertile patches in arid systems and describes previous research on the effects of ecosystem engineers in arid environments. Chapter 2 examines how fertile patch creation by a well-studied arid zone engineer, the ant, varies between ant species and landforms, demonstrating that while we can generalise about the effects of ant nests on water flow and nutrient levels, differences in soil type, nest density and ant species across sites are likely to moderate these effects. Chapters 3, 4 and 5 compare the engineering effects of reintroduced native species, the Greater bilby (Macrotis lagotis) and Burrowing bettong (Bettongia lesueur) with the Sand goanna (Varanus gouldii) and the invasive European rabbit (Oryctolagus cuniculus) across three Australian landforms, revealing differences between reintroduced and invasive engineers at the both patch and landscape scales. At the landscape scale, differences in pit densities and resource concentration in foraging pits between landforms suggest that fertile patch development is not uniform. Chapter 6 examines whether the structure of the pit or the presence of litter per se explains observed increases in plant germination in foraging pits, and compares temperature and soil moisture conditions in bilby and bettong pits to the soil surface. Chapter 7 is a review that builds upon previous chapters, published and unpublished literature of the potential for reintroductions of ecosystem engineers to restore ecosystem function in degraded environments. Chapter 8 summarises the research presented in this thesis and its implications, and suggests directions for future work.

Marine ecosystems are increasingly being subject to human impact from diverse recreational and commercial activities, not necessarily restricted to those of a marine nature. This has significant implications for biodiversity. The large dog cockle, Tucetona laticostata, once occurred live in Rangitoto Channel, Hauraki Gulf, although this species no longer appears to occur there, most likely as a consequence of repeated dredging and channel excavation and continued siltation. Tucetona laticostata still occurs in a few isolated pockets of sea bed throughout Hauraki Gulf, particularly off Otata Island, part of the Noises complex of islands, where it resides partially buried in shell and rock gravels in shallow water (to 15 metres depth). The shells of T. laticostata collect in large post-mortem deposits in an area ramping from the sea bed off southwestern Otata Island. These mounds differ significantly in structural complexity from those of adjacent, extensively fragmented shell and rock gravels. Using the mounds of T. laticostata shell as a proxy for structural complexity, in order to appraise the effect of complexity on benthic-invertebrate diversity, the sea bed off southwestern Otata Island was sampled quarterly at two depths and in both T. laticostata shell mounds and adjacent extensively fragmented shell and rock gravels. These data were complemented with those from additional surveys around Otata Island, and off eastern Motutapu Island to determine the distribution and composition of benthic-invertebrate community assemblages throughout the region, and from concurrent surveys throughout the Waitemata Harbour and inner Hauraki Gulf to determine the current distribution of T. laticostata in this region. The number of benthic invertebrate species and individuals within T. laticostata habitat almost always was higher than that occurring within extensively fragmented shell- and rock gravel habitat, with densities to 142,385 individuals m-2 encountered. Temporal and spatial variations in benthic community structure also are reported for the two habitats, T. laticostata-based shells and extensively fragmented shell- and rock gravels. The numbers of species were higher amongst samples collected off the southwestern and eastern sides of Otata Island than elsewhere around this island, or of eastern Motutapu Island. Of the 351 species reported from all Otata and Motutapu Island samples combined, 73% of them occurred off southwestern Otata Island, 30% of which were found exclusively within T. laticostata shell habitat, and 10.5% within extensively fragmented shell and rock gravel habitat. The sea bed off southwestern Otata Island is regularly, seasonally dredged by recreational scallop fishers, in addition to being a popular small-vessel anchorage site. Both of these activities, dredging and anchorage, stand to reduce substratum complexity by fragmentation and dispersal of the valves of T. laticostata. Given the unique benthic invertebrates reported from T. laticostata shell deposits reported from southwestern Otata Island, any activity that damages the shells of this species, regardless of whether they are live or dead, is likely to result in loss of biodiversity. Admittedly, many of species identified as major contributors to differences in benthic invertebrate assemblages between T. laticostata shell-based habitats and those of extensively fragmented shell and rock gravels are not particularly charismatic or large, but each likely plays a role in local food webs and/or sediment and water column chemistry. It was not the intention of this research to determine the effects of anthropogenic disturbances like dredging or vessel anchorage on benthic-invertebrate communities off southwestern Otata Island. However, given the reported differences in species diversity within the structurally complex substratum provided by T. laticostata, conservation of biogenic reef-forming species like it might be a prudent, precautionary measure to take.

The modifications animals make to their environments can be critical to species diversity. Red squirrels (Tamiasciurus hudsonicus) create large piles of conifer cone debris (middens) and are hypothesized to function as a keystone species due to positive associations between middens and other vertebrate species. We assessed vegetation and landscape structure at middens with a resident red squirrel for varying consistencies over the prior 5 years and surveyed mammals and birds at the community and population level. After accounting for vegetation and landscape characteristics, red squirrel-created resources positively influenced species richness of medium and large mammals and ground foraging birds, abundance of deer mice (Peromyscus maniculatus) and ground foraging birds, and activity of cliff chipmunks (Tamias dorsalis). Increased knowledge of the interaction strength of the red squirrel with its environment may be used to inform decisions in forest management and restoration and offer insight on the conservation value of larderhoarding mammals.

abstract: As much as 40% of the world's human population relies on rivers which originate on the Qinghai-Tibetan Plateau (QTP) (Xu et al. 2009, Immerzeel et al. 2010). However, the high alpine grasslands where these rivers emanate are at a crossroads. Fed by seasonal monsoon rains and glacial runoff, these rivers' frequent flooding contributes to massive losses of life and property downstream (Varis et al. 2012). Additionally, upstream grasslands, which regulate the flow of these rivers, are considered to be deteriorating (Harris 2010). This thesis examines the regional vulnerability of these rivers and highlights the impacts of several policy responses, finding that both climate change and grassland degradation pose significant challenges to Asia's water security. Additionally, I suggest that many of the responses elicited by policy makers to meet these challenges have failed. One of these policies has been the poisoning of a small, endemic, burrowing mammal and keystone species, the plateau pika (Ochotona curzoniae) (Smith and Foggin 1999). Contrary to their putative classification as a pest (Fan et al. 1999), I show that the plateau pika is instead an ecosystem engineer that actively increases the infiltration rate of water on the QTP with concomitant benefits to both local ecosystems and downstream hydrological processes. / Dissertation/Thesis / M.S. Biology 2012

Orientador: Prof. Dr. Guilherme Henrique Pereira Filho / Dissertação (mestrado) - Universidade Federal do ABC, Programa de Pós-Graduação em Evolução e Diversidade, 2016. / Engenheiros Ecossistemicos (EE) modulam a disponibilidade de recursos no ambiente. Este e o caso dos rodolitos (i.e. gnodulos de vida livreh compostos por mais de 50% de algas calcarias incrustantes que formam bancos em fundos inconsolidados) que promovem habitats para diversos organismos. Por outro lado, muitos desses organismos modulam condicoes para o crescimento de rodolitos. Esta interacao resulta na criacao de redes altamente dependentes de EEs. O peixe Malacanthus plumieri move rodolitos para a construcao de montes. Esta movimentacao altera o estado fisico de nodulos e, possivelmente, contribua para a diferenciacao das comunidades associadas. Portanto, avaliamos se as comunidades de macrofauna vagil (> 500¿Êm) presentes nos montes sao afetadas de forma positiva, negativa ou neutra em relacao a area controle (i.e. sem interferencia do M. plumieri). Alem disso, procuramos identificar as principais variaveis fisicas, que sao alteradas pelo peixe, que predizem a estrutura da macrofauna associada aos rodolitos. Para tanto, amostramos no Arquipelago de Fernando de Noronha 30 rodolitos na area controle e 30 nos montes. Cada nodulo foi individualmente inserido em sacolas de nylon com malha de 500¿Êm. Os epibiontes foram removidos e identificados. Mensuramos volume e diametros de cada rodolito e quantificamos o peso seco das macroalgas e do sedimento. Observamos que rodolitos da area controle apresentaram maior quantidade de macroalgas epifitas e sedimentos aprisionados quando comparados com rodolitos presentes nos montes (t=-0,04, p<0,001 e t=-0,54, p <0,001; respectivamente). A bioturbacao causada por M. plumieri altera as condicoes para o estabelecimento da macroalgas, reduzindo as taxas de sedimentos aprisionados. Como resultado, ha uma reducao na abundancia, riqueza e densidade de macroinvertebrados (t=-7,42, p<0,001; t=-7,92, p<0,001; t=-2,84, p<0,01; respectivamente). Portanto, M. plumieri afeta indiretamente a comunidade de macroinvertebrados vageis. Em contraste com outros estudos, o efeito da coexistencia destes EEs parece ser antagonico (i.e. positivo para a abundancia e riqueza de ictiofauna e negativo para macroalgas e macroinvertebrados). Assim, mudancas estruturais no sistema, embora promovidas pelo mesmo EE, influenciam comunidades de diferentes maneiras e sao dependentes da escala dos organismos em questao. / Ecosystem engineers (EE), such as rhodoliths, control the availability of resources in the environment, providing habitats for other organisms, as other algae and many animals. Some of these organisms are able to modulate the environmental conditions where rhodoliths grow, thus forming a highly dependent network. The sand tilefish Malacanthus plumieri, for example, moves rhodoliths to build mounds over their burrows. This behavior changes the physical medium in which the nodules are, and therefore, their associated communities¿ structure and composition. We assessed the effect of communities associated with mounds in comparison to non-mounded areas (i.e. rhodolith areas without the interference of the tilefish). Additionally, we seek to identify the main physical variables changed by the action of M. plumieri in predicting the structure of the macrofauna associated with rhodoliths. Rhodoliths from non-mounded areas showed higher amounts of epiphytic macroalgae and sediment trapped on rhodoliths than mounds (t=-0.04, p<0.001 and t=-0.54, p<0.001, respectively). Bioturbation caused by M. plumieri alters the conditions for macroalgae establishment, reducing the trapped sediment. As a result, there is also a reduction in the abundance, richness and density of macroinvertebrates (t=-7.42, p<0.001, t=-7.92, p<0.001, t=-2.84, p<0.01 respectively). The ecosystem role of M. plumieri indirectly affects the vagile macrofauna community. In contrast to other studies, the coexistence of these EEs (i.e. rhodoliths and tilefishes) seems to be antagonistic (i.e. positive for the abundance and richness of ichthyofauna and negative for macroalgae and macroinvertebrates). Thus, structural changes in the system, although promoted by the same EE, influence communities in different ways depending on the scale of the organisms in question.