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Università Degli Studi Di Cagliari PHD DEGREE Earth and Environmental

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Università Degli Studi Di Cagliari PHD DEGREE Earth and Environmental 1 2 3 4 5 6 7 8 Università degli Studi di Cagliari 9 10 11 12 PHD DEGREE 13 Earth and Environmental Sciences and Technologies (EEST) 14 Cycle XXXIII 15 16 17 Distribution, floristic and ecological characterization of Sardinian Gymnospermae communities 18 19 Botanica Ambientale e Applicata (BIO/03) 20 (Environmental and Applied Botany) 21 22 23 24 PhD Student: Dr. Giacomo Calvia 25 Coordinator of the PhD Programme Prof. Giorgio Ghiglieri 26 Supervisor Prof. Gianluigi Bacchetta 27 Co- Supervisors Prof. Emmanuele Farris 28 Prof. Giuseppe Fenu 29 30 31 Final exam. Academic Year 2019 - 2020 32 Thesis defence: July 2021 33 1 34 2 35 GENERAL INTRODUCTION 5 36 1.1 General information about Gymnosperms 6 37 1.2 Study area: the island of Sardinia 8 38 1.3 Conifer species in Sardinia 9 39 1.4 The selected conifers and their features 10 40 1.5 Research objectives and structure of this thesis 14 41 1.6 References of introduction 16 42 CHAPTER 1: Classification of the Sardinian pine woods 21 43 1. Introduction 23 44 2. Material and methods 24 45 2.1 Study area 24 46 2.2 Pine species of interest 24 47 2.3 Data collection 25 48 2.4 Data preparation and analyses 26 49 3. Results 26 50 4. Discussion 36 51 5. Conclusions 37 52 6. References 39 53 7. Appendixes 42 54 CHAPTER 2: Temporal increase in the extent of pine stands in Sardinia 52 55 1. Introduction 54 56 2. Material and methods 55 57 2.1 Study area 55 58 2.2 Pine species 55 59 2.3 Current distribution of native pine habitats of Sardinia 56 60 2.4 Diachronic analysis of the distribution of the pine species in Sardinia 56 61 2.5 Collection and interpretation of the pine-related toponyms of Sardinia 57 62 3. Results 57 63 3.1 Current distribution of the pine species and habitats in Sardinia 57 64 3.2: Diachronic analysis of the distribution of natural pine woods in Sardinia (1954-2016) 59 65 3.3 Pine-related toponyms 60 66 4. Discussion 60 67 4.1 Distribution of the three pine species in Sardinia: an update 60 68 4.2 Pine wood changes between the 20th and 21st centuries reflect the land-use change 62 69 5. Conclusions 63 70 6. References 64 3 71 7. Appendices 68 72 CHAPTER 3:From real to potential distribution: the case of study of Taxus baccata L. in Sardinia 73 73 1. Introduction 75 74 2. Material and methods 76 75 2.1 Study area 76 76 2.2 Target species 77 77 2.3 Data collection and analysis 77 78 3. Results 78 79 3.1 Yew distribution in Sardinia 78 80 3.2 Yew potential distribution in Sardinia 83 81 4. Discussion 84 82 5. References 86 83 6. Appendices 91 84 CHAPTER 4: Ecological and anthropic factors affecting dead wood and regeneration of Sardinian yew 85 (Taxus baccata L.) stands: implications for conservation 92 86 1. Introduction 94 87 2. Material and method 96 88 2.1 Study area 96 89 2.2 Data collection 97 90 2.3 Statistical analysis 97 91 3. Results 98 92 4. Discussion 102 93 4.1 Implications for conservation 103 94 5. Conclusions 104 95 6. References 104 96 7. Appendices 108 97 SUMMARY 111 98 1.1 General conclusions 111 99 1.2 Final considerations 111 100 4 101 General introduction 102 5 103 1.1 General information about Gymnosperms 104 Gymnosperms represent an important part of the plant world. Despite being only 0.29% of the entire plant 105 kingdom, compared to a total 89.4% of Angiosperms (Crepet & Niklas 2009), Gymnosperms are widespread 106 in many areas of the world, sometimes dominating in density, cover and biomass over the other taxa (Farjon 107 2017). They are present in all continents, excepted Antarctica, and their distribution has often been enlarged 108 by human activities (Eckenwlder 2009; Farjon 2017). 109 Conflicting studies about the origin of Gymnosperms and Angiosperms had been carried out, but it is still not 110 well clear which were the dynamics related to speciation and separation of the various groups (Farjon 2008; 111 Christenhusz et al. 2011). The Gymnosperms had been considered for a long time a distinctive group of plants 112 and many unresolved problems and debates get involved the different groups (Earle 2021). Currently, 113 molecular analyses have not removed issues as the origin of a presumed common ancestor within the different 114 Gymnosperms. Besides, it is not resolved the doubt about the existence of a common ancestor shared both by 115 the Gymnosperms and the flowering plants or Angiosperms (Eckenwalder 2009). According to Armstrong & 116 Brasier (2005), Angiosperms would have been evolved from an advanced group of gymnosperms, but still 117 controversial relationships occur between the two groups of seed plants. Palynological data report a first 118 appearance of Angiosperms during the Early Cretaceous (Hughes & McDougall 1987) same era where the first 119 record of the genus Pinus is known (Ryberg et al. 2012), while the earliest seed plants are known from the 120 early Devonian (Hill 2005). 121 All the Gymnosperms are woody, mostly trees, shrubs, and vines (Earle 2021). Their name derives from the 122 ancient Greek word γυμνόσπερμος (γυμνός, gymnós, "naked" and σπέρμα, spérma, "seed"), meaning "naked 123 seeds". That is due to the condition of their ovules, which are not enclosed in a capsule (ovary) but are often 124 naked (Farjon 2017). 125 The Gymnosperms are separated in two divisions, Gnetophyta and Pinophyta (Earle 2021): to Gnetophyta 126 Bessey 1907 belong the three sub-classes Cycadidae Persoon ex Bercht. et J. Presl, Ginkgoidae Engl. and 127 Gnetidae Pax ex Pranti. 128 Cycadidae is a subclass represented by the only order Cycadales Persoon ex Bercht. et J. Presl, in which are 129 included three families: Cycadaceae L., Stangeriaceae L.A.S.Johnson, and Zamiaceae Horianow, for a total 130 extent of about 290 species, mostly resembling palms (Earle 2021). All the species grow at tropical and 131 subtropical latitudes, in all the continents excepted Europe (Jones 1993). 132 The subclass Ginkgoidae includes the order Ginkgoales Gorozh., with a single family, Ginkgoaceae Engl., 133 consisting of the monotypic Ginkgo biloba L., which is considered a living fossil and is today very rare in the 134 wild, being known in some small areas of south-western China, in the provinces of Guizhou and Zhejiang 135 (Gong et al. 2008; Earle 2021). 136 The subclass Gnetidae has three orders: Ephedrales Dumort., Gnetales Blume ex von Martius, and 137 Welwitschiales Skottsburg ex Reveal (Christenhusz et al. 2011). In the order Ephedrales is included one single 138 family (Ephedraceae Dumort.) and one genus (Ephedra L.) composed by 61 taxa (Earle 2021). Many species 139 grow in the Eurasia, others in Americas and Africa, normally in arid and semiarid environments from the sea 140 level up to 5000 m a.s.l. in the Himalayan and Andean Chains (Price 1996). The order Gnetales is represented 6 141 by the only family Gnetaceae Blume, with one genus (Gnetum L.) and 44 species (Earle 2021). Many of the 142 species belonging to this family are vines and grow in south-eastern Asia (Carlquist 1996), while a smaller 143 number of species is known in South America and in western Africa (Price 1996). Finally, the order 144 Welwitschiales is represented by the monotypic family Wellwitschiaceae Caruel, with Welwitschia mirabilis 145 Hooker, one of the strangest plants in the world, having only two leaves that constantly grow during all its life. 146 It is limited to a small area of south-western Africa (Namib desert; Eller et al. 1983). 147 The division Pinophyta Raveal includes the only subclass Pinidae Cronquist, Takhtajan et Zimmermann, which 148 is divided in three orders: Pinales Gorozh., Araucariales Gorozh., and Cupressales Link. To the order Pinales 149 belongs the family Pinaceae Spreng. ex Rudolphi, with 232 species included in 11 genera (Farjon 2017). 150 Almost all the entities of this family grow in the northern hemisphere, both in temperate and sub-arctic 151 environments, as well as in sub-tropical areas (Farjon 2017). On the other hand, the order Araucariales includes 152 two families, Araucariaceae Henkel et W. Hochstetter and Podocarpaceae Endl., that are mostly distributed in 153 the southern hemisphere, while in the northern one they grow only on tropical regions (Farjon 2017): the 154 Araucariaceae are divided in three genera (Araucaria Jussieu, Agathis Salisb., and Wollemia Jones, Hill et 155 Allen) and about 40 species. The family Podocarpaceae is larger, including 18 genera and 174 species (Farjon 156 2017). The order Cupressales comprises three different families (Sciadopityaceae Luerss., Cupressaceae Rich. 157 ex Bartling and Taxaceae Gray). The family Sciadopityaceae is monotypic, represented only by Sciadopitys 158 verticillata (Thunberg) Siebold et Zuccarini, a tree growing in Japan (Tsukada 1963). The Cupressaceae family 159 is the richest within conifers in genera, being 27, which include a total 143 species and subspecies 160 (Eckenwalder 2009). It is also the most widespread conifer family, being known in all the continents excepted 161 Antarctica (Earle 2021). Molecular analyses showed the former family Taxodiaceae C.N.Page was to include 162 within Cupressaceae (Brunsfeld et al. 1994). Finally, the family Taxaceae is represented by 6 genera and 28 163 species, mostly diffused in the boreal hemisphere (Earle 2021). 164 This PhD research is focused on a specifical group of Gymnosperms, i.e. conifers (Pinophyta). Conifers 165 characterise many natural environments, mostly in the northern hemisphere of the world (Eckenwlder 2009; 166 Debreczy & Racz 2011; Farjon 2017).
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