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Proved to Be Appendages Are Arranged In The Winteraceaeof the Old World. I. Pseudowintera and Drimys — Morphology and taxonomy W. Vink Summary is southwest The primitive woody Angiospermfamily Winteraceae centred in the Pacific, has an outpost in Madagascar (1 species), and a section ofDrimys (4 species) in the New World. Pseudowintera is restricted to New Zealand, the Old World section Tasmannia of Drimys extends from the Philippines to Tasmania. Since A. C. Smith’s review of the family in 1943 about ten times as much material has become available the the section Tasmannia is it and it appears that taxonomy of far more complicated than seemed to be in New and attention its 1943. This induced me to make a field study in Guinea to pay special to morphology in the hope offinding additional criteria. On this basis the Australian species of Drimys could be better defined but the characters of the New Guinean taxa(Smith: 29 species) proved to be highly unstable. It was found that in the section Tasmannia the stomata are occluded by waxy plugs. The structure ofthe is in section In flower inflorescence described and compared with that Drimys. the two zones are distinguish- ed. In the lower zone the floral appendages are arranged in (sub)opposite pairs, in the higher zone they are distance from focus of arranged according to the space available at the greatest possible a the floral apex which I termed ‘activity centre’; this centre often does not coincide with the ‘topographical apex’. The tilt of determination in ofthe floral apex appears to influence the arrangements as well. Shifting factors relation the available locations the floral is assumed role. to on apex to play an important As New Guinean The for the specific delimitation,the material appeared to be the crux. very heteroge- in it neous material did not show correlated discontinuities characters, which made impossible to reach a division into The of the is described with species. variability resulting very complex species Drimys piperita of the aid a non-taxonomic subdivision, the units ofwich are called ‘entities’. Field studies revealed that local often but the discontinuities between such local populations are very stable, morphological populat- ions is are obscured by material from other localities. It supposed that vegetativepropagation as observed in the combined with incidental be of in such situation. field, cross-breeding, may importance establishing a In Australia besides D. piperita four other species can be distinguished. The sections of Drimys are maintained,although Ehrendorfer c.s. and Smith proposed raising them to generic rank. In Pseudowintera within is described the arrangement the inflorescence spiral, not decussate as earlier, and a terminal flower is Minute hairs bracts and the mode of of the present. on young leaves, opening calyx fact cupule, and details of the lower leaf surface are used as a new set of specific characters. The that about ofthe show combinations of characters is be caused twenty per cent specimens varying specific thoughtto by hybridisation. Contents Introduction and acknowledgements 226 The genus Pseudowintera Introduction 227 Morphology I. Twig 227 2. Lower leaf surface 227 3. Inflorescence 228 4. Flower 228 226 BLUMEA VOL. XVIII. No. 2, 1970 Fruit 5. 231 6. Seed 231 7. Conclusions 233 and Sporogenesis embryology 233 Specific delimitation 233 Specific descriptions 235 Taxonomy 235 The genus Drimys Morphology I. Introduction 244 2. Twig 244. 3. Leaf 246 4. Occlusion of the stomata 246 3. Inflorescence 248 6. Flower 259 7. Fruit 263 8. Seed 264 Ontogeny of the flower 266 D. 266 I. Sect. Tasmannia: piperita 2. Sect. Tasmannia: D. purpurascens, D. stipitata, D. insipida 274 3. Sect. Tasmannia: D. lanceolata 274 4. Sect. Drimys 277 Diagram of the flower 277 floral Comparison of arrangements of appendages 278 Reproduction 281 Specific delimitation in Sect. Tasmannia 283 I. New Guinea 283 ia. Mt Wilhelm 284 lb. Mt Wilhelmina complex 287 Mt ic. Ambua c.s 289 id. Entity 'heteromera' c.s 295 ie. Use of characters 299 if. Conclusion 301 2. Philippines to Moluccas, and Flores 301 3. Australia 301 Phytochemistry 301 Taxonomy 302 Fossils 351 Excluded from the Winteraceae 351 Literature 352 Index 353 Introduction and acknowledgements Since A. C. Smith published his 'Taxonomic notes on the Old World species of Winteraceae' (J. Arn. Arb. 24, 1943, 119 —164) no further taxonomic work has been the the done on this family. Such work is more important as Smith had no access to in last material in the European herbaria because of the war; moreover, the two decades the collections have increased enormously, enlarging our possibilities for research. To illustrate this for the available point: 1943 review were 25 New Guinean collections, whereas had from for the present work I 480 collections that area at may disposal. this the Pseudowintera and the section Tasmannia of the In paper genus genus Drimys are have from treated. I refrained including the American section Drimys as this has been A. fully revised by C. Smith (I.e., pp. 1—33). of the characters of the will be in future when the A discussion family given a paper remaining genera have been treated. W. Vink: The Winteraceae of the Old World. I. 227 This study could not have been made without the generosity of the directors of the following herbaria, Arnold who kindly put their collections at my disposal: Adelaide, Arboretum, Berlin, Bishop Museum, British Museum of Natural History, Bogor, Canberra, Christchurch, Copenhagen, Florence, Kew, Lae, Leiden, Manilla, Melbourne,Paris, Rancho Santa Ana, Singapore, Stockholm, Sydney, Utrecht, Washing- ton, Wellington, and Zurich. Netherlands Foundation for Advancement of Research financed The the Tropical WOTRO an explo- the Doma in New make extensive ration of Peaks area Guinea, where I was able to collections. I am much indebted to the Director of the Departmentof Scientific and Industrial Research at Christ- New Dr. W. C. church for information concerningthe Zealand localities, and to van Heel and Dr. Kalkman Mrs. for criticism. To C. den Hartog, I am most grateful for the considerable improvements ofthe English text. THE GENUS PSEUDOWINTERA INTRODUCTION review of the taxonomicand nomenclatural of the Pseudowintera For a history genus I refer to the excellent introductions and by J. E. Dandy, J. Bot. 71 (1933) 119—121 A. C. from Smith, J. Arn. Arb. 24(1943) 154—155. For the complications arising the typification of Drimys, see p. 237, 303. MORPHOLOGY The of Pseudowintera bear normal leaves I. Twig. twigs (long shoots) only, cataphylls are of leaves and not formed. The terminalbud consists young is usually very small; it often disintegrates, an axillary bud taking over as leader. The short shoot, forming an axillary inflorescence, may sometimes produce apically a vegetative long shoot. The bracts are then often slightly larger and longer persistent than the shoot in normal inflorescence. The result is a normal lateral bearing, however, in its basal in their axils. part cataphylls (enlarged bracts) with flowers, fruits, or scars of these this it in Although is a rare phenomenon, appears regularly some specimens (fig. 4 d). Nast (1944) and Sampson (1963) report the development of vegetative shoots in the axils of bracts, replacing the secondary inflorescences (see below). For of a description the structure of the vegetative shoot apex, see Gifford (1950) 603, 605. 2. Lower leaf surface. The cuticle of the lower leaf surface consists of two layers. The inner one, next to the outer wall of the epidermis cells, is homogeneous and has often distinct plasmodesma-like structures (described by Sampson as 'invaginations, apparently of the wall into the cuticle'). Overlying this layer is a non-homogeneous, white-coloured which layer is alveolar according to Bailey & Nast (1944b). In P. colorata this alveolar layer carries the in fine tubercles. Although not actually observed, presence of waxes or on the alveolar layer is deduced from the disappearance of the white colour when applying gentle heat. The white colour can be restored by boiling and drying the leaf. It is assumed that the wax melts when heat is applied, replacing the air or the crystalline structure of the wax in the interstices of the alveolar layer. After cooling the white colour does not return as this is caused by reflection on the interstices filled with air or crystalline wax. the Boiling will remove the wax and subsequent drying admits the air into interstices, thus restoring the white colour. Over the subsidiary cells and the guard cells the homogeneous layer thins out, while the alveolar the thickness. The front of the stomata layer at same places increases in cavity is completely occluded by alveolar material. Whether the occlusion is provided with 228 BLUMEA VOL. XVIII. No. 2, 1970 dotted: Fig. 1. Schematic cross-sections through lower leaf surfaces of Pseudowintera; homogeneous P. P. cuticular layer; hatched: alveolar material. a. P. axillaris. b. colorata. c. traversii. narrow canals or with interstices larger than those in the alveolar layer over the epidermis cells, could not be established. The three species of Pseudowintera show different combinations of the situations de- scribed above andof theposition ofthe stomata hi relation to the surface of the epidermis, as follows: axillaris: P. alveolar layer absent over epidermis cells; surface of the occlusion not sunken below the general surface of the epidermis (fig. ia). colorata: alveolar cells and P. layer present over epidermis provided with minute tubercles; surface of the occlusion not sunken below the general surface of the epidermis (fig. ib). alveolar of the white P. traversii: layer present over part epidermis cells, forming blotches or a discontinuous layer, not provided with tubercles; surface of the occlusion distinctly sunken below the general surface of the epidermis (fig. ic). Inflorescence. The inflorescence consists 3.
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