Umschlagbild des Buches: Richard Bolli - Revision of the <i>Genus Sambucus</i>

Richard Bolli:

Revision of the Genus Sambucus

1994. II, 256 pages, 14x22cm, 510 g
Sprache: English

(Dissertationes Botanicae, Band 223)

ISBN 978-3-443-64135-1, paperback

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BibTeX Datei

Schlagworte

elderberry • Holunder • sureau • sambucus nigra • Caprifoliaceae

Inhalte

Synopsis nach oben ↑

More than 100 names for species, subspecies and varieties of Sambucus, elderberry, have been created so far but no more than about 30 of these are usually taken into account in Floras. This is due to the work of F. Graf von Schwerin, the first to give comprehensive treat­ents of the genus in 1909 and 1920. As he grew the majority of the species and many cultivars in his garden in northern Germany, he was able to observe the great variability of pubescence and carpel merosity, formerly regarded as important diagnostic characters, and consequently he reduced the number of species recognized by him to 28.

However, as typical of his time, he still overestimated the systematic value of vegetative characters and fruit colour variants, maintaining or even newly describing species of very doubtful status.

Still, we are confronted with an intriguing and puzzling variability and heterogeneity within the genus, despite its homogeneous and characteristic general appearance. Whereas branching architecture, leaves, inflorescences and flowers are extremely similar in all species, life forms and fruit colours differ notably between popu­lations in definite geographic ranges. The close relationships within the genus are further reflected by a striking ecological similarity. All Sambucus species prefer moist and nutrient-rich habitats. Only two taxa are resistant to seasonal drought. In their natural habitats, but especially in secondary stations, which are invaded by all species, the stout perennials or many-branched shrubs or trees grow very fast. They may either colonize wide areas as pioneers or become integrated into the shrub layers of various, mainly mesophytic, woodland ecosystems.

Moreover, the uniqueness of the genus gives good reasons for classifying Sambucus in a family all by itself, Sambucaceae. The only approach to unravelling these complex variation patterns is to search for new characters with the modern methods from different biological disciplines such as morphology, cytology, phytochemistry, and biogeography. The results of this multidisciplinary approach will permit to unravel the systematic relationships within the genus and put forward an appropriate taxonomy.

While describing variability patterns of each of the species, an attempt was made to keep their world-wide connections in mind. This taxonomic revision was undertaken from a global point of view and the conclusion was reached that many closely similar taxa must be placed together in a small number of species, rather than putting them in a large array of hardly definable, allegedly distinct species. I hope that the Sambucus taxonomy here proposed here will be useful for both field and herbarium botanists.

Beyond the interest of botanists, elders are popular and widely known plants. Leaf and flower extracts, and occasionally fruits, are used in herbal medicine, for treating colds and coughs, by almost all cultures of both the Old and the New World where Sambucus occurs. The fruits are widely used as raw material for preparing a delicious syrup, jam or wine. Yet, the question of toxicity of some species is under debate in popular herb books. Thus, it is understandable that some Sambucus species have successfully followed man’s cultivation.

Table of contents nach oben ↑

Acknowledgements 4
Introduction 5
Haterial and Hethods 7
Field Studies 7
Herbarium Studies 7
Morphological Methods and Measurements 8
General Part
Growth Habit, Root and Stem 11
Growth Habit 11
Rhizomatous Perennial Herbs 11
The Woody Species 11
The Root 17
The Stem 17
Hood Anatomy 20
Generic Description 20
Taxonomic and Ecological Implications 21
The Leaves 23
Stipules and Extrafloral Nectaries 32
Ecology of the Extrafloral Nectaries 33
Inflorescence and Flower 35
The Inflorescence 35
The Flower 38
Variation in the Number of Floral Organs 41
Pollen 42
The Ovary 43
Pollination Biology and Breeding System 44
Hypothesis 44
Observations and Pollination Experiments 44
Pollen Viability and Pollen Tube Growth 48
Conclusions 51
Hybrids and Cultivars 51
Fruit and Seed 52
Number of Loculi 53
Fertility 55
Dispersal and Distribution 57
Natural Dispersal 57
Dispersal by Man 59
Range Margins and Outposts 61
General Distribution and Comparison with other Genera 61
Life Cycle 64
Germination 64
The Seedling 66
Establishment of the Plantlet 66
The Fertile Phase 66
What Age Can an Elder Tree Reach? 67
Death 68
The Habitat 69
Chemical Constituents, Toxicity and Use 73
Toxicity: Are there Poisonous Species in Sambucus? 73
The Cyanogenic Glycoside Sambunigrin as a Chemical Deterrent 75
Ecology of Cyanogenesis 76
Medical and Culinary Use 77
Fruit Anthocyanins 79
Material and Methods 79
Thin Layer Chromatography (TLC) 82
Results and Discussion 82
Conclusions 84
High Performance Liquid Chromatography (HPLC) 85
Results and Discussion 85
In Which Way Is Fruit Colour Determined? 94
Biosynthetic Pathway and Evolutionary Significance 95
Hypothetical Biosynthetic Pathway of Sambucus Anthocyanins 96
Genetic Background of Sambucus Anthocyanins: A Hypothesis 97
Evolutionary Interpretation of Sambucus Anthocyanin Data 99
Karyosystenatics 101
Chromosome Numbers 101
Chromosome Morphology 104
Discussion
Visualization of Evolution in the Genus 105
Evolutionary Significance of some Important Characters 105
Growth Habit 105
Inflorescence 106
Flower 106
Pollen Exine Sculpturing 107
Fruit Anthocyanins 108
Karyology 109
Character Combinations and Conflicting Characters 109
Earth History and Biogeography 113
An Old World Herbaceous Perennial as Ancestor of Sambucus 115
Place and Time of Origin 115
Scenario 1: Late Cretaceous Origin 116
Scenario 2: Oligocene Origin 119
Systematic Part
The Family Sambucaceae 125
Where to Place Sambucus? 125
Sambucus, Viburnum and Adoxa 126
The Genus Sambucus 129
Sambucus L 129
Infrageneric Classifications 130
Taxonomic Species Concept 132
Subspecies and Species 133
Variety and Continuous Variation 133
Note 1: How to Collect Sambucus Specimens 134
Note 2: Citation of Types 134
Key to the Species and Subspecies 135
1. Sambucus ebulus L 140
S. ebulus sap. ebulus 143
S. ebulus ssp. africana (Engler) R.Bolli stat.nov 144
2. Sambucus nightiana Wallich ex Night et Arnott 146
3. Sambucus adnata DC 149
4. Sambucus gaudichaudiana DC 151
5. Sambucus australasica (Lindley) Fritsch 155
6. Sambucus javanica Blume 158
7. Sambucus nigra L 161
S. nigra ssp. nigra 162
S. nigra ssp. maderensis (Lowe) R.Bolli stat. et comb. nov 164
S. nigra ssp. palmensis (Link) R.Bolli stat. et comb. nov 167
S. nigra ssp. canadensis (L.) R.Bolli stat. et comb. nov 168
S. nigra ssp. cerulea (Rafin.) R.Bolli stat. et comb. nov 174
S. nigra ssp. peruviana (H.B.K.) R.Bolli stat. et comb. nov. 177
8. Sambucus australis Cham, et Schlecht 181
9. Sambucus racemosa L 187
Preliminary Remarks on Sambucus racemosa L 187
S. racemosa var.racemosa 195
S. racemosa var.melanocarpa (Gray) McMinn 195
Names of Taxa of Uncertain Application 198
Summary 199
Zusammenfassung 201
References 205
Appendix
Synoptic Key to the Species of Sambucus 223
Index of Taxa 224
Index of Plates 227