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Stygobites) in England and Wales DOCTORAL THESIS An investigation into the distribution of obligate groundwater animals (stygobites) in England and Wales Weitowitz, Damiano Award date: 2017 General rights Copyright and moral rights for the publications made accessible in the public portal are retained by the authors and/or other copyright owners and it is a condition of accessing publications that users recognise and abide by the legal requirements associated with these rights. • Users may download and print one copy of any publication from the public portal for the purpose of private study or research. • You may not further distribute the material or use it for any profit-making activity or commercial gain • You may freely distribute the URL identifying the publication in the public portal ? Take down policy If you believe that this document breaches copyright please contact us providing details, and we will remove access to the work immediately and investigate your claim. Download date: 09. Oct. 2021 An investigation into the distribution of obligate groundwater animals (stygobites) in England and Wales N. kochianus captured from the Chalk in England, UK. Damiano C Weitowitz BSc, MSc A thesis submitted in partial fulfilment of the requirements for the degree of PhD Department of Life Sciences University of Roehampton 2016 II Abstract Aquifers and their biota (stygobites) are some of the least well known ecosystems. Previous research indicates geology is an important control of stygobites and the primary thesis aim was therefore to examine how geology shapes the distributions of stygobites across different spatial scales, considering high lithological detail. The food web functionality of groundwater ecosystems is also rarely explored and the secondary thesis aim was therefore to examine the trophic effects of stygobites in experimental groundwater microcosms. The new habitat typology shows that 11 higher resolution geo- habitats are characterised by significantly different hydrochemistry and a heterogeneous distribution of high-quality habitat patches. Furthermore, the habitat scoring system developed based on the variability of influential abiotic parameters shows that overall geo-habitat quality varies considerably, with karstic geo-habitats (e.g. Chalk) having a higher quality than most porous and fractured geo-habitats. Major parts of England and Wales are covered by poor quality habitats, probably limiting dispersal. Testing the new typology on species distributions shows that biodiverse and / or abundant communities occur in all geo-habitats. Karstic aquifers generally form the best habitats with the highest stygobite species diversity and frequency of occurrence. However, some fractured aquifers (e.g. Igneous Rock) are also significant habitats for stygobites, while more geological detail needs to be considered to explain stygobite communities in other fractured aquifers (e.g. Mudstones & Siltstones). However, many species are not distributed throughout entire connected aquifers, and in individual sites copepod biomass is not explained by geology, showing that other environmental controls (glacial history, site characteristics) may supersede geological controls. The grazing experiments show 2 Abstract that stygobites significantly increase protozoan abundance and morphotype diversity, and alter microbial community structure, indicating they may modulate the ecosystem services provided by these groups. Overall, this thesis enhances our knowledge of the factors governing the distribution and functionality of groundwater ecosystems. 3 III Contents Title Page………………………………………………………………………………………………………………………p.1 II Abstract……………………………………………………………………………………………………………………..p.2 III Contents……………………………………………………………………………………………………………………p.4 Tables………………………………………………………………………………………………………………………..p.9 Figures…………………………………………………………………………………………………………………………p.11 Appendices………………………………………………………………………………………………………………….p.15 IV Acknowledgments…………………………………………………………………………………………………. p.18 1. General Introduction……………………………………………………………………………………………p.20 1.1 Summary of thesis aims……………………………………………………………………………………p.20 1.2 Groundwater characteristics…………………………………………………………………………….p.22 1.3 Conservation status of groundwater ecosystems……………………………………………..p.29 1.4 Geology – A control across multiple scales……………………………………………………….p.30 1.5 Current habitat typologies……………………………………………………………………………….p.32 1.6 A global sampling bias………………………………………………………………………………………p.37 1.7 Groundwater controls on faunal biomass…………………………………………………………p.41 1.8 Geology and associated heterogeneity as stygobite controls……………………………p.44 1.9 Surface water and groundwater food webs……………………………………………………..p.47 2. Defining geo-habitats for groundwater ecosystem assessment (submitted to Hydrogeology journal)……………………………………………………………………….p.53 2.1 Abstract……………………………………………………………………………………………………………p.54 2.2 Introduction……………………………………………………………………………………………………..p.55 2.3 Methodology……………………………………………………………………………………………………p.58 4 Contents 2.3.1 Assessing geo-habitat distribution………………………………………………………..p.59 Determining geo-habitat categories………………………………………….p.59 Assigning geological units to geo-habitats…………………………………p.62 2.3.2 Assessing geo-habitat characteristics……………………………………………………p.65 Data Collection………………………………………………………………………….p.65 Comparing Geo-Habitat Characteristics…………………………………….p.66 2.3.3 Evaluation of Geo-Habitat Quality………………………………………………………..p.67 Identification of thresholds and additional parameters…………….p.67 Calculation………………………………………………………………………………..p.68 2.4 Results……………………………………………………………………………………………………………..p.70 2.4.1 Distribution of geo-habitats………………………………………………………………….p.70 2.4.2 Geo-habitat characteristics…………………………………………………………………..p.72 Hydrogeology……………………………………………………………………………p.74 Hydrochemistry…………………………………………………………………………p.77 2.4.3 Geo-habitat quality………………………………………………………………………………p.79 2.5 Discussion………………………………………………………………………………………………………..p.84 2.5.1 Assessing the new typology………………………………………………………………….p.84 2.5.2 Geo-habitat characteristics and quality………………………………………………..p.86 2.5.3 Distribution of geo-habitats………………………………………………………………….p.91 2.6 Conclusion……………………………………………………………………………………………………….p.94 2.7 Acknowledgments……………………………………………………………………………………………p.95 2.8 References……………………………………………………………………………………………………….p.96 2.9 Appendices…………………………………………………………………………………………………….p.106 3. The groundwater ecology of poorly sampled geologies in England and Wales………………………………………………………………………………………………..p.111 3.1 Abstract………………………………………………………………………………………………………….p.112 3.2 Introduction…………………………………………………………………………………………………..p.114 3.3 Methodology………………………………………………………………………………………………….p.118 3.3.1 Distribution of fauna…………………………………………………………………………..p.118 5 Contents Selection of geologies and sampling areas………………………………p.118 Field sampling…………………………………………………………………………p.120 Laboratory sorting and identification………………………………………p.122 Assessing species distributions……………………………………………….p.123 3.3.2 Calculating individual and total biomass…………………………………………….p.125 Individual biomass…………………………………………………………………..p.125 Total biomass in geologies………………………………………………………p.125 3.3.3 Assessing environmental predictors of copepod biomass and…………….p.125 abundance 3.4 Results……………………………………………………………………………………………………………p.128 3.4.1 Faunal distributions in geologies…………………………………………………………p.128 3.4.2 Biomass trends in geologies……………………………………………………………….p.137 3.4.3 Individual copepod biomass and abundance………………………………………p.139 Local site effects……………………………………………………………………..p.139 Other environmental effects…………………………………………………..p.140 3.5 Discussion………………………………………………………………………………………………………p.144 3.5.1 Stygobite distributions in the four geologies………………………………………p.144 3.5.2 Overall biomass pattern……………………………………………………………………..p.148 3.5.3 Influences on copepod biomass and abundance…………………………………p.151 3.6 Conclusion……………………………………………………………………………………………………..p.154 3.7 Acknowledgments………………………………………………………………………………………….p.156 3.8 References……………………………………………………………………………………………………..p.157 3.9 Appendices…………………………………………………………………………………………………….p.170 4. Geological controls on stygobite distributions in England and Wales………………….p.175 4.1 Abstract………………………………………………………………………………………………………….p.176 4.2 Introduction…………………………………………………………………………………………………..p.177 4.3 Methodology………………………………………………………………………………………………….p.181 4.3.1 National-scale assessment of stygobite distributions………………………….p.181 Data Sources…………………………………………………………………………..p.181 6 Contents Geo-habitat suitability for stygobites………………………………………p.184 Geological controls of stygobite distributions…………………………p.185 4.3.2 Stygobite habitat chemistry………………………………………………………………..p.186 Data Sources…………………………………………………………………………..p.186 Water chemistry ranges of stygobites…………………………………….p.187 4.4 Results……………………………………………………………………………………………………………p.188 4.4.1 Geo-habitat suitability for stygobites………………………………………………….p.188 4.4.2 Geological controls of stygobite distributions…………………………………….p.189 4.4.3 Species habitat chemistry differences…………………………………………………p.202 4.5 Discussion………………………………………………………………………………………………………p.205 4.5.1 Geo-habitat suitability for stygobites………………………………………………….p.205 4.5.2 Geological controls on stygobite distributions…………………………………….p.209 4.5.3 Stygobite water chemistry ranges………………………………………………………p.216 4.6 Conclusions……………………………………………………………………………………………………p.218 4.7 Acknowledgments………………………………………………………………………………………….p.220 4.8 References……………………………………………………………………………………………………..p.221 4.9 Appendices…………………………………………………………………………………………………….p.233 5. 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