Hughes, Oliver Thomas Wesley (2018)Orchid – relationships, propagation of terrestrial and epiphytic orchids from seed. Doctoral thesis (PhD), Manchester Metropolitan University.

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Orchid – Mycorrhiza relationships, propagation of terrestrial and epiphytic orchids from seed

Oliver Thomas Wesley Hughes

A thesis submitted in partial fulfilment of the requirements of Manchester Metropolitan University for the degree of Doctor of Philosophy

Division of Biology and Conservation Ecology

School of Science and the Environment

In Collaboration with the North of England Zoological Society and the University of Manchester

2018

1

Contents

Abstract ...... 14

Acknowledgements ...... 16

Chapter 1...... 20

General Introduction ...... 20

References ...... 29

Chapter 2...... 38

Symbiotic germination of the terrestrial orchid media in vitro is not dependent on fungal sympatry ...... 38

Abstract ...... 39

Keywords Terrestrial orchid germination · Fungal compatibility orchids · Mycorrhizal Fungi · · Symbiotic propagation ...... 39

Introduction...... 39

Materials and Methods ...... 40

Study species ...... 40

Experimental design ...... 42

Results ...... 43

Discussion ...... 45

Acknowledgements ...... 50

References ...... 50

Chapter 3...... 55

Assessing fungal compatibility in the germination and development of four terrestrial orchids of the subtribe Orchidinae ...... 55

Abstract ...... 56

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Keywords • Mycorrhizae • Fungal compatiibility • Germination • Conservation ...... 56

Introduction...... 56

Materials and Methods ...... 59

Organismal details ...... 59

Experimental procedure ...... 61

Results ...... 64

Anacamptis morio ...... 66

Dactylorhiza incarnata ...... 67

Serapias bergonii ...... 72

Serapias parviflora ...... 74

Discussion ...... 75

Species specific responses ...... 75

Experimental variables ...... 77

Fungal specificity and compatibility ...... 79

Acknowledgements ...... 81

References ...... 82

Appendix ...... 90

Chapter 4...... 92

Symbiotic culture enhances in vitro germination and development of two European terrestrial orchids; laxiflora and ... 92

Abstract ...... 93

Keywords Orchidaceae • Germination • Terrestrial • Mycorrhizae • Conservation ...... 93

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Introduction...... 94

Materials and methods ...... 97

Results ...... 101

Initial germination and development ...... 101

Further development and acclimatisation ...... 105

Discussion ...... 109

Acknowledgements ...... 114

References ...... 114

Chapter 5...... 120

The effects of substrate rugosity and complexity on in vitro germination and development in the European terrestrial orchids; Anacamptis morio and purpurella ...... 120

Abstract ...... 121

Keywords Orchidaceae • Germination • Terrestrial • Mycorrhizae • Rugosity • Substrate complexity ...... 121

Introduction...... 122

Materials and methods ...... 124

Organismal details ...... 124

Experimental procedure ...... 125

Results ...... 129

Germination and development ...... 129

Statistical analysis ...... 135

Rugosity treatments ...... 135

Negative control (treatments) ...... 136

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Substrate complexity ...... 137

Discussion ...... 138

Acknowledgements ...... 144

References ...... 144

Chapter 6...... 162

Asymbiotic and symbiotic germination of five epiphytic orchids from the subtribe - advantage for initial symbiotic germination 162

Abstract ...... 163

Keywords Orchidaceae • Mycorrhizae • Germination • Propagation • Conservation ...... 164

Introduction...... 164

Materials and Methods ...... 168

Organismal details ...... 168

Experimental procedure ...... 169

Results ...... 173

Masdevallia melanoxantha...... 174

Germination ...... 174

Development ...... 175

Masdevellia menatoi ...... 176

Germination ...... 176

Development ...... 177

Masdevallia patchicutzae ...... 179

Germination ...... 179

Development ...... 179

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Pleurothallopsis microptera ...... 179

Germination ...... 179

Development ...... 180

Restrepia contorta ...... 181

Germination and development ...... 181

Stelis sp. Peru ...... 181

Germination and development ...... 181

Discussion ...... 186

Acknowledgements ...... 191

References ...... 192

Chapter 7...... 203

A method for using microcosms for the ex situ study of tripartite symbioses between ectomycorrhizal , fungi and myco-heterotrophic plants ...... 203

Abstract ...... 204

Keywords Orchidaceae • Mycorrhizae • Myco-heterotrophy • Microcosm • Symbiotic culture ...... 204

Introduction...... 204

Materials and methods ...... 206

Organismal details ...... 206

Experimental procedure ...... 207

Results ...... 209

Discussion ...... 212

Acknowledgements ...... 215

6

References ...... 215

Appendix ...... 222

Chapter 8...... 225

General Discussion ...... 225

References ...... 237

7

List of Figures

Chapter 1.……………………………………………………………………………….. 21

General Introduction………………………………………………………………….. 21

Chapter 2………………………………………………………………………………... 38

Symbiotic germination of the terrestrial orchid Microtis media is not dependent on fungal sympatry……………………………………………………... 38

Figure 1. Number of Microtis media protocorms on each of the fungal isolates……………………………………………………………………………... 44 Figure 2. Stages of development in Microtis media ………………...... 45

Figure 3. Microtis media plants with Rhizoctonia solani……………………… 46

Chapter 3………………………………………………………………………………... 55

Assessing fungal compatibility in the germination and development of four terrestrial orchids of the subtribe Orchidinae……………………………… 55

Figure 1. Stages of orchid germination used to assess development………. 65

Figure 2. Germination of Anacamptis morio, , Serapias bergonii and Serapias parviflora on each treatment……………….. 68

Figure 3. Stages of development in Anacamptis morio, Dactylorhiza incarnata, Serapias bergonii……………………………………… 70

Figure 4. Development of Anacamptis morio on each treatment…………… 71

Figure 5. Development of Dactylorhiza incarnata on each treatment…….... 72

Figure 6. Development of Serapias bergonii on each treatment……..…….. 73

Figure 7. Development of Sera