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Download This Document Mortimer, E.J. (2016) Ecotypic variation in 'Lotus corniculatus L.' and implications for grassland restoration: interaction of ecotypes with soil type and management, in relation to herbivory. PhD thesis. Bath Spa University. ResearchSPAce http://researchspace.bathspa.ac.uk/ Your access and use of this document is based on your acceptance of the ResearchSPAce Metadata and Data Policies, as well as applicable law:- https://researchspace.bathspa.ac.uk/policies.html Unless you accept the terms of these Policies in full, you do not have permission to download this document. This cover sheet may not be removed from the document. Please scroll down to view the document. Ecotypic variation in Lotus corniculatus L. and implications for grassland restoration: Interaction of ecotypes with soil type and management, in relation to herbivory Erica Jane Mortimer A thesis submitted in partial fulfilment of the requirements of Bath Spa University for the degree of Doctor of Philosophy School of Society, Enterprise & Environment, Bath Spa University 2016 This copy has been supplied on the understanding that it is copyright material and that no quotation from the thesis may be published without proper acknowledgement. I have exercised reasonable care to ensure that my thesis is original, and does not to the best of my knowledge break any UK law or infringe any third party’s copyright or other intellectual property right. Word count: 37515 Ancillary Data: 23116 (Contents/glossary/acknowledgements: 3311; tables/figures/captions: 10438; references: 9367) Appendices: 16417 2 ABSTRACT This research assesses the importance of using ecologically-similar rather than geographically-local seed in grassland restoration projects, with particular reference to herbivorous invertebrates, including pollinators. Seed from Lotus corniculatus L. (bird’s-foot trefoil) populations at nine sites across south-west England were collected to represent ecotypes potentially adapted to a range of soil types (calcareous loam, neutral loam and calcareous sand (referred to as ‘sand’)) and management regimes (grazed, cut with aftermath grazing (referred to as ‘cut’) and unmanaged). From October 2011 the ecotypes were planted within three different treatment soils (calcareous loam, neutral loam and calcareous sand (“sand”), and two management treatments (grazed [simulated] or unmanaged [neither cut or grazed]). Differences in plant morphology and phenology under these treatments were recorded at four-weekly intervals and immediately prior to harvest on 16th July 2012. Fresh and dry biomass were recorded and leaf- nitrogen and leaf-hydrogen cyanide (HCN) levels determined. Treatments were maintained and plants grown for a further 15 months [after harvest]. Bee preference for ecotypes grown under treatment combinations was also recorded during peak flowering periods of 2012 and 2013. Data were tested using Kruskall-wallis and ANOVA. A Generalized Linear Mixed Model (GLMM) was built to test all ecotype and treatment differences including interactions. A separate Non-linear Mixed Effects model (NLME) was created to investigate spatial autocorrelation between ecotype sites. The standard chosen in the models was Cockey Down (a calcareous loam, grazed ecotype) grown in matching treatments. The phenotypic traits retained were most pronounced in ecotypes from home- sites of sand soil type and cut management, which were considered to be the more stressed environments of the study, requiring rapid adaptation. By harvest, ecotypes from sand home-sites produced significantly greater number of stems per plant, greater leaflet number per main stem and lower 3 HCN. Model results for sand ecotypes additionally identified delayed seed pod formation, increased hirsuteness and higher leaf-HCN compared to the standard. Significant differences found within ecotypes from cut home-sites included fewer stems per plant, fewer leaflets per main stem, more seeds per pod, greater leaf-HCN and shorter time to first flower. The model also found this ecotype to be less hirsute with fewer seed pods (in unmanaged treatment) than the standard. Ecotypic traits shown in plants from the less stressed home-sites included calcareous loam ecotypes having two clear flowering peaks in both years and highest leaf-HCN, and unmanaged ecotypes having lower leaf-HCN. Three significant interactions indicated additive character factor effects: neutral loam ecotypes grown in neutral loam treatment soil had earlier pod formation than the standard; and, sand ecotypes grown in sand treatment soil and unmanaged ecotypes receiving unmanaged treatment had greater flower number (over both years) than all other ecotypes, treatments and combinations. Results from the bee ecotype preference study showed no preference for ecotypes geographically close to the test foraging area. However significant differences were shown by bees in terms of ecotype preference, with avoidance of plants containing highest leaf-HCN. Plants grown in calcareous soil treatment were preferred which suggests nectar of plants are of most value to bees when grown in optimum [for L. corniculatus growth] soil. Eotypic differences in herbivory defence [leaf-HCN and hirsuteness induced by home-site soil and management], would be of importance to receptor site invertebrate herbivores/pollinators. Pollinators could also find difficulties with the ecotypic differences in flowering asynchronicity. Both home-site soil type and management could also influence the viability of the plant population through reducing fecundity. Delayed seed pod formation in sand ecotypes (compared to the standard) indicates an adaptation to summer temperatures or low-nutrient availability. Calcareous loam ecotypes lack such adaptation and if introduced to a sand sites and could fail due to 4 poorly timed germination, deep seed burial [from shifting sand] or poorly allocated energy. Cut ecotypes also produced significantly fewer seed pods (than the standard) with significantly greater seed numbers per pod suggesting an energy allocation adaptation to produce fewer, larger pods before defoliation rather than continuous pod formation throughout the season, a potentially critical adaptation for seed return. Findings from this study are of national relevance, and Natural England should adopt new recommendations on seed provenance in agri-environment schemes. Instead of recommending strictly geographically local seed, the management regime (particularly details of intensity and timing of management operations) should ideally be similar between the donor and receptor sites. Soil types, especially pH and clay/organic matter content should also be matched as far as possible as these were the greatest limiting factors within this study. Suitable donor sites may be local sites of similar habitat. If no such sites are available then recommendations from this study should be followed in seeking suitable sowing material. If seed suppliers are used, then they should provide greater detail on donor site conditions to aid land managers. 5 CONTENTS ABSTRACT 3 CONTENTS 6 Table of Figures 11 Table of Tables 15 GLOSSARY 17 ACKNOWLEDGEMENTS 19 1 INTRODUCTION 20 1.1 Biodiversity Loss and Habitat Restoration 20 1.2 Restoration Ecology and Seed Provenance 23 1.3 Wider Ecology of Habitat Restoration 25 1.4 Need for Further Research 26 2 AIM AND OBJECTIVES 27 2.1 Aim 27 2.2 Objectives 27 3 OVERVIEW OF METHODS USED 29 3.1 Ecotypes 29 3.2 Soil Treatment (Study 1) 30 3.3 Management Treatment (Study 2) 30 3.4 Monitoring and Harvest 30 3.5 Post-Harvest Monitoring 31 3.6 Bee Interaction (Study 3) 32 4 STUDY SPECIES LOTUS CORNICULATUS L. 33 4.1 Natural History 33 4.2 Agriculture 34 4.3 Conservation 35 4.4 Plant Defences 36 5 ECOTYPE SITES – INITIAL COLLECTIONS 39 5.1 Ecotypic Variation in Seed 39 5.2 Factors in Choice 40 5.3 Ecotype Site Choices 40 6 5.4 Sites with Permissions 41 5.5 Seed Collection Methodology 46 6 ECOTYPE SOILS (DONOR SITES) 49 6.1 Soil Type Introduction 49 6.2 Ecotype Soil Collection and Analysis Methodology 51 6.2.1 Soil Collection 51 6.2.2 Soil Chemical and Physical Analysis 51 6.2.3 Data Analysis 54 6.3 Ecotype Soil Results 54 6.3.1 All Ecotype Soils 54 6.3.2 Chosen Ecotype Soils 58 6.4 Ecotype Soil Discussion 60 7 ECOTYPE SEED (Donor Site) 63 7.1 Seed Preparation and Sowing Methodology 63 7.1.1 Seed Preparation 63 7.1.2 Seed Sowing 64 7.1.3 Data Analysis 66 7.2 Seed Weight and Germination Results 66 7.3 Seed Weight and Germination Discussion 69 8 FINAL ECOTYPE CHOICE 70 8.1 Overview of Ecotype Selection 70 8.2 Geographical Distance between Chosen Ecotype Sites 73 9 SOIL TYPE TREATMENTS (RECEPTOR SITES) 75 9.1 Soil Types in Grassland Restoration 75 9.2 Treatment Soil Initial Choice 76 9.3 Treatment Soil Collections and Analysis Methodology 76 9.3.1 Pre-Collection Decisions 76 9.3.2 Mole Hill Soil Properties 77 9.3.3 Soil Collection 77 9.3.4 Soil Chemical and Physical Analysis and Final Treatment Choice 78 9.3.5 Data Analysis 79 9.4 Treatment Soil Results 80 9.4.1 Results of All Potential Treatment Soils 80 9.4.2 Further Results of Chosen Treatment Soils 84 9.5 Treatment Soil Discussion 86 9.6 Treatment Soil Final Choice 86 9.6.1 Win Green – Calcareous loam 87 7 9.6.2 Avis Meadow – Neutral loam 87 9.6.3 Woolacombe Warren – Sand 87 10 MANAGEMENT REGIME TREATMENT 88 10.1 Management of Grasslands 88 10.1.1 Intensive Management 88 10.1.2 No Management 88 10.1.3 Cut with Aftermath Grazing 89 10.1.4 Extensive Grazing 90 10.2 Species Adaptation to Management 91 10.3 Grassland Restoration 91 10.4 Management
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