Chapter Two: the Ecological Character of Bodmin Moor
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University of Plymouth PEARL https://pearl.plymouth.ac.uk 04 University of Plymouth Research Theses 01 Research Theses Main Collection 2018 Taxonomy, biogeography and ecology of Andean tardigrades at different spatial scales Ramsay, Balbina http://hdl.handle.net/10026.1/12178 University of Plymouth All content in PEARL is protected by copyright law. Author manuscripts are made available in accordance with publisher policies. Please cite only the published version using the details provided on the item record or document. In the absence of an open licence (e.g. Creative Commons), permissions for further reuse of content should be sought from the publisher or author. Taxonomy, biogeography and ecology of Andean tardigrades at different spatial scales by BALBINA P. L. RAMSAY A thesis submitted to the University of Plymouth in partial fulfilment for the degree of DOCTOR OF PHILOSOPHY School of Biological and Marine Sciences Faculty of Science and Engineering January 2018 ii Copyright Statement This copy of the thesis has been supplied on condition that anyone who consults it is understood to recognise that its copyright rests with its author and that no quotation from the thesis and no information derived from it may be published without the author's prior consent. iii iv Acknowledgements I thank Dr Dave Bilton and Dr Simon Rundle (my PhD advisors) for their support and advice during the course of the doctorate. Dr Paul Ramsay, University of Plymouth, provided the original idea, organised the field work, helped to collect the samples and critically reviewed the manuscript. Nigel Marley assisted greatly with tardigrade processing and taxonomy which was a crucial part of this work. I also wish to thank Roy Moate, Pete Bond and Glenn Harper from the Electron Microscopy Centre at the University of Plymouth. I gratefully acknowledge the assistance of the people from Mantanay Valley, Peru for granting access to their Polylepis forest. The Ecuadorian Ministry of the Environment provided permits to collect and borrow the sample material used in this study: Scientific Investigation Authorisation Patent No. 10-2010-FLO-DPAP-MA and Patent No. 009- 2015-FAU-DPAP-MA. The QCA Herbarium and the QZNE Museum at the Pontificia Universidad Católica del Ecuador helped with the permits. I also acknowledge, with gratitude, the advice and assistance of David Suarez Duque and Segundo Chimbolema for their help in the field which were important for this work. Claire Rampage and Magdalena Fiedler translated some taxonomic papers from Russian and Polish to English, respectively, which were important in Chapter 2. v vi Author's declaration At no time during the registration for the degree of Doctor of Philosophy has the author been registered for any other University award without prior agreement of the Graduate Committee. Relevant scientific seminars and conferences were attended at which work was presented; external institutions were visited for consultation purposes and several papers prepared for publication. Presentations Given: Ramsay, B.P.L and Ramsay, P.M. (2016) Diversidad de tardígrados en un bosque de Polylepis en el norte de Ecuador. IV Congreso International de Ecología y Conservación de Bosques de Polylepis, San Salvador de Jujuy, Argentina 5-9 September 2016. Ramsay, B.P.L (2011) La biogeografía de tardígrados y su estudio. Symposium presentation to the Department of Biological Sciences, Pontificia Universidad Católica del Ecuador, Quito, Ecuador, August 2011. Ramsay, B.P.L (2011) Biogeography, ecology and taxonomy of high-altitude Andean tardigrades. Marine Biology and Ecology Research Centre Seminar Series, University of Plymouth, United Kingdom. Ramsay, B.P.L and Marley, N.J. (2009) Tardigrade communities from the high-altitude forest to grassland transition in the Cordillera de Vilcanota, Peru. XI International Symposium on Tardigrada, Tübingen, Germany, July 2009. Conferences Attended: XIII International Symposium Tardigrada, Modena, Italy, 23-26 June 2015 – Attendance XII International Symposium on Tardigrada, Vila Nova de Gaia, Portugal, 23-26 July 2012 – Attendance Word count of main body of thesis: 31,405 Signed: Date: 22 May 2018 vii viii Balbina Pérez Lara Ramsay: Taxonomy, biogeography and ecology of Andean tardigrades at different spatial scales. Abstract Micrometazoans are animals smaller than 2 mm. Their biogeography is poorly understood, and tardigrades provide a tractable phylum for exploring distribution patterns at a variety of scales. Polylepis forest habitat offers considerable advantages for making tardigrade comparisons across a wide range of scales in the Andes. This thesis aims to improve identifications of tardigrades with a character matrix approach, to assess the relative importance of habitat and bryophyte host on tardigrades, to describe the fine-scale spatial structure of tardigrade assemblages, and to estimate the sampling effort required for a reliable estimate of tardigrade diversity within Polylepis forest. Samples of bryophytes and lichens were collected from Polylepis forest and neighbouring habitats, and the tardigrades extracted and identified, mostly to operational taxonomic units. Some new species were discovered during the course of this work; one is described here. Abundance, diversity and composition of tardigrade samples were compared quantitatively. The thesis presents the first example of a character matrix for a tardigrade genus, bringing together information for the genus Isohypsibius from many different sources and describing suites of characters for each species. It will facilitate identification within the genus in future. Tardigrade assemblage data were highly variable within the samples, with empty samples dominating one study. Analysis of one forest site indicated that at least 50 samples would be needed to characterise the tardigrade diversity there. Although both were important, habitat- scale effects were more influential on tardigrade abundance, diversity and composition than host-scale effects. In both cases, microenvironmental and resource filters are the likely mechanisms driving these differences. Based on the results, recommendations are made for expanding such research into broader geographical scales: standardising sample volume, replicate sampling across hosts on the forest floor, recognising the importance of habitat-scale effects when selecting study sites, and the development of character matrices for tardigrade genera. ix x List of Contents 1. Spatial scale and the biogeography of Tardigrada: Polylepis forests in the Andes as a model system ............................................ 1 Aims of the thesis .............................................................................. 12 Outline of the thesis ........................................................................... 12 2. Development of a character matrix to describe a new species of Tardigrada, Isohypsibius saulrodgersi sp. nov. (Eutardigrada, Isohypsibiidae), from Ecuador ..............................................15 Introduction ...................................................................................... 15 Materials and methods ....................................................................... 16 Results .............................................................................................. 18 Discussion ......................................................................................... 33 Conclusions ....................................................................................... 35 3. The effect of microhabitat on the distribution of tardigrades at high- altitude forest in the Peruvian Andes ....................................37 Introduction ...................................................................................... 37 Materials and methods ....................................................................... 38 Results .............................................................................................. 40 Discussion ......................................................................................... 48 4. The structure of tardigrade communities at the landscape scale: the influence of habitat and host ...............................................53 Introduction ...................................................................................... 53 Methods ............................................................................................ 55 Results .............................................................................................. 56 Discussion ......................................................................................... 64 Conclusions and recommendations ...................................................... 69 5. The structure of tardigrade communities at fine spatial scales in an Andean Polylepis forest .......................................................71 Introduction ...................................................................................... 71 Methods ............................................................................................ 73 Results .............................................................................................. 76 Discussion ......................................................................................... 83 Conclusions ....................................................................................... 88 6. General discussion ..............................................................89 7. References .........................................................................97