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Systematic Wood Anatomy of Myodocarpus, Delarbrea, and Pseudosciadium (Araliaceae)

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Systematic wood anatomy of Myodocarpus, Delarbrea, and Pseudosciadium (Araliaceae) Alexei A. OSKOLSKI Botanical Museum, V. L. Komarov Botanical Institute of the Russian Academy of Sciences, Prof. Popov Str. 2, 197376 St. Petersburg, Russia. Porter P. LOWRY II Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. [email protected] Laboratoire de Phanerogamie, Museum national d'Histoire naturelle, 16 rue Buffon, 75005 Paris, France. [email protected] Hans Georg RICHTER Bundesforschungsanstalt für Forst- und Holzwirtschaft, Leuschnerstrasse 91, 21031 Hamburg, Germany. ABSTRACT Wood anatomical features examined in five species of Myodocarpus, four spe­ cies of Delarbrea, and the single member of Pseudosciadium (Araliaceae) using light and scanning electron microscopy support the hypotheses that they are a closely related, monophyletic group, and that they form an ancient lineage that has survived and diversified on the island of New Caledonia, where all except two of the 17 species belonging to these genera are endemic. Delarbrea and Pseudosciadium have similar wood sttucture; the wood of Myodocarpus is distinctly more primitive (sensu BAILEY). These genera differ markedly in wood structure from other Araliaceae in the presence of apotta- cheal (diffuse and diffuse-in-aggregates) axial parenchyma, and wood features KEYWORDS Myodocarpus, do not suppott earlier suggestions that Myodocarpus is intermediate between Delarbrea, Araliaceae and Apiaceae. Within Myodocarpus two groups of species can be Pseudosciadiu m, distinguished on wood characters, which correspond to the species with Araliaceae, wood anatomy, simple vs. pinnately compound leaves. The wide-ranging D. paradoxa subsp. vasicentric tracheids, paradoxa is the only taxon studied with numerous vasicentric tracheids, often phylogeny, New Caledonia. regarded as an adaptation to watet stress. ser. 3 • 1997 • 19(1) 61 Oskolski A A, Lowry II P.P. & Richter H.G. RÉSUMÉ Les caractères anatomiques du bois ont été examinés pour cinq espèces de Myodocarpus, quatre espèces de Delarbrea, et l'unique espèce de Pseudosciadium (Araliaceae) aux microscopes photonique et électronique à balayage. Les résultats obtenus confirment que ces genres sont proches, constituent un groupe monophylétique et forment une lignée ancienne qui a survécu et s'est diversifiée en Nouvelle-Calédonie, où 15 des 17 espèces de ces genres sont endémiques. La structure du bois de Delarbrea et Pseudosciadium est semblable ; celle de Myodocarpus est bien plus primitive (sensu BAILEY). Ces trois genres ont une structure anatomique du bois nette­ ment différente de celle des autres Araliaceae en raison de la présence d'un parenchyme axial apotrachéal (diffus à diffus-agrégé). L'ensemble des carac­ tères observés n'est pas compatible avec l'idée que Myodocarpus occupe une MOTS CLES Myodocarpus, position intermédiaire entre les Araliaceae et les Apiaceae. Chez les Delarbrea, Myodocarpus deux groupes d'espèces peuvent être distingués par l'anatomie Pseudosciadium, du bois : celles à feuilles simples et celles à feuilles composées. Delarbrea Araliaceae, anatomie du bois, paradoxa subsp. paradoxa, à large répartition, est le seul taxon étudié à possé­ trachéides vasicentriques, der de nombreuses trachéides vasicentriques, caractère souvent considéré phylogénie, Nouvelle-Calédonie. comme une adaptation au stress hydrique. INTRODUCTION floral vasculature is also relatively primitive (EYDE & TSENG 1971). However, Myodocarpus, Myodocarpus, Delarbrea, and Pseudosciadium Delarbrea, and Pseudosciadium differ from all form a closely related group of genera centred in other Araliaceae by the presence of distinctive New Caledonia (LOWRY 1986a, 1986b). All ten secretory oil ducts in the fruits (EYDE & TSENG species of Myodocarpus and the single species of 1971; LOWRY 1986a, 1986b), which appear to Pseudosciadium are endemic to this island, situa­ tepresent a derived character (synapomorphy). ted in the southwest Pacific. Moreover, four of Moreover, the apotracheal axial parenchyma the six species of Delarbrea are likewise restricted found in the wood of the first two genera is also to New Caledonia; D. paradoxa Vieill. subsp. unknown among other members of the family paradoxa also extends through Vanuatu and the (RODRIGUEZ 1957; OSKOLSKI 1994; but see Solomon Islands to the Moluccan and Lesser below regarding Pseudosciadium). Sunda Islands, and D. michieana (F. v. Muell.) Myodocarpus stands out by being the only genus F. v. Muell., previously placed in the monotypic of Araliaceae with a dry, laterally compressed, genus Porospermum F. v. Muell., is endemic to schizocarpic fruit with a central carpophore, Queensland, Australia (LOWRY 1986a, 1986b). whose overall morphology is at least superficially The plants of this alliance are monocaulous or similar to the fruits of many Apiaceae (by sparsely branched treelets or trees ranging from contrast, the fruits of Delarbrea and ca. 1.5 to 20 m in height, with leaves that are Pseudosciadium are terete drupes). On this basis, densely clustered at the branch ends, and whose BAUMANN (1946) regarded Myodocarpus as a pos­ petiole bases are clasping, as in many other sible link between Araliaceae and its traditional Araliaceae. The three genera share a number of sister group, Apiaceae (see also THORNE 1973), other characters, including similar inflorescence which together are widely recognized as compri­ structure and organization, articulated pedicels, sing the order Araliales. More recently, however, an isomerous androecium, and a bicarpellate a number of studies have questioned such an gynoecium, although each of these features also intermediate position for the genus (RODRIGUEZ occurs individually elsewhere in the family. Their 1957, 1971; EYDE & TSENG 1971; LOWRY 62 ADANSONIA, sér. 3 • 1997 • 19(1) Wood anatomy of Araliaceae 1986a, 1986b; OSKOLSKI 1994; PLUNKETT 1994; of the three genera under consideration here that PLUNKETT et al. 1996). LOWRY (1986a) points was included in his study) is part of an ancient out that the fruits of all three genera share a group from which both Araliaceae and Apiaceae single basic anatomical plan, with each tissue evolved. present in the fleshy fruits of Delarbrea and Structural features of wood offer a useful tool Pseudosciadium having a direct homologue in for improving our understanding of the systema­ those of Myodocarpus, including the characteris­ tic position of Myodocarpus, Delarbrea, and tic oil ducts. Pseudosciadium within Araliaceae, as well as the The close relationship between Delarbrea and relationships among these three genera. Myodocarpus was first pointed out by VlElLLARD Previously published data on the wood structure (1865). BENTHAM (1867), and later HARMS of these taxa are scanty, and hence insufficient to (1894-97), HUTCHINSON (1967), and TSENG & develop any conclusive interpretations. Only Hoo (1982) placed these genera in the tribe four species of Myodocarpus have been studied Aralieae Benth., whereas VlGUIER (1906) and previously [M. simplicifolius Brongn. & Gris TAKHTAJAN (1987) treated them as members of a (KRIBS 1937; OSKOLSKI 1994); M. pinnatus segregate tribe Myodocarpeae (erroneously refer­ Brongn. & Gris (SARLIN 1954); M. fraxinifolius red to as Myodocarpineae by VlGUIER). The sys­ Brongn. & Gris and M. involucratus Dubard & tematic placement of Pseudosciadium has a Vig. (RODRIGUEZ 1957); Myodocarpus sp. somewhat more complex history. When BAILLON (RECORD & HESS 1944; METCALFE & CHALK (1878, 1879) first described Pseudosciadium 1950)], while only a single member of Delarbrea balansae, he indicated that it was closely related had been examined [D. paradoxa subsp. paradoxa to both Delarbrea and Myodocarpus, and further (OSKOLSKI 1994)]. suggested that it was intermediate between them. In later treatments of Araliaceae (HARMS This study, which is part of a general survey of 1894-97; VIGUIER 1906, 1925; HUTCHINSON wood anatomy throughout the family (OSKOLSKI 1967), however, Pseudosciadium was separated 1994, 1995, 1996; OSKOLSKI & LOWRY in from both genera, and included in the tribe prep.), examines the wood structure of a much Mackinlayeae, which was rigidly (and artificially) larger sample of Myodocarpus, Delarbrea, and defined by valvate, clawed petals. The close rela­ Pseudosciadium, and considers the results with tionship of Pseudosciadium to Delarbrea and regard to the systematic relationships of the Myodocarpus initially suggested by BAILLON was group. The conclusions from such systematic confirmed when oil ducts were observed in the analyses of wood structural features represent an fruits of Pseudosciadium by LOWRY (1986a, important contribution to an overall understan­ 1986b), who concluded on the basis of this and ding of the family, and will assist in preparing other characters that these three genera comprise further revisions of Araliaceae for the Flore de la a monophyletic group. He further suggested that Nouvelle-Calédonie (LOWRY 1986a, 1986b, in they represent the relicts of an ancient aralia- prep.) and other regions in the Pacific (LOWRY ceous lineage forming part of a floristic ensemble 1987, 1988, 1989, 1990, in prep.; LOWRY et al. that was able to survive in the relatively equable 1989). Moreover, this type of study provides climates of New Caledonia, but was in large part additional morphological information for eliminated from Australasia and elsewhere as a ongoing comparative phylogenetic analyses based result of changing climatic conditions in largely on molecular data (PLUNKETT 1994;
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    Index References to accepted names in bold-faced print, to synonyms in upright print, to illustrations in italics. A A. quaternata, 258 Annesorhiza, 101 Absolmsia, 333 A. sect. Tonduzia, 259 Anodendron, 302 Acanthopanax, 441 Alstonieae, 257 Anomalluma, 342 Acerates, 370 Alyxia, 276 Anomopanax, 84 Aciphylla, 95 A. concatenata, 276 Anomotassa, 381 Acokanthera, 280 Alyxieae, 274 Anthanotis, 370 Acomosperma, 392 Amalocalyx, 306 Anthocleista, 493 Acrocoryne, 382 Amarella, 488 Anthoclitandra, 264 Acronema, 96 Ambelania, 267 Anthriscus, 102 Actinolema, 92 Amblyopetalum, 386 A. caucalis, 102 A. macrolema, 92 Ammi, 98 A. caucalis var. caucalis, 102 Actinotus, 82 A. majus, 14 A. caucalis var. gymnocarpa, 102 Acustelma, 317 Ammodaucus, 99 A. kotschyi, 102 Adelostemma, 377 Ammoides, 99 A. sylvestris, 18, 102 Adelphacme, 520 Ammoselinum, 99 Antiostelma, 333 Adenium, 285 Ampelamus, 377 Antonia, 520 Adenolisianthus, 497 Amphidetes, 391 Antoniaceae, 511 Adenosciadium, 96 Amphineurion, 302 Antonieae, 520 Aegokeras, 96 Amphistelma, 380 Antoniinae, 520 Aegopodium, 96 Amphorella, 388 Apegia, 344 Aenigma, 397 Amsonia, 271 Aphanopleura, 102 Aethusa, 97 Amsonieae, 271 Aphanostelma, 386 Aframmi, 137, 158 Anagallidium, 489 Aphanostylis, 264 Afrocarum, 107 Anakasia, 436 Apiaceae, 9 Afroligusticum, 97 Anatropanthus, 329 Apiastrum, 103 Afrosciadium, 97 Ancylobothrys, 263 Apioideae, 95 Afrosison, 170 Andrewsia, 486 Apiopetalum, 82 Aganonerion, 305 Andriana, 99 A. glabratum, 83 Aganosma, 306 Anechites, 278 A. velutinum, 83 A. sect. Amphineurion, 302 Anemotrochus, 387 Apium, 103 Agasyllis, 97 Anethum, 99 Apocinae, 207 Agathotes, 489 Angadenia, 294 Apocynaceae, 207 Agrocharis, 97 Angelica, 100 Apocyneae, 301 Aidomene, 370 A. czernaevia, 18 Apocynoideae, 281 Alafia, 286 A. decurrens, 14 Apocynoids, 281 Albovia, 170 Anginon, 100 Apocynum, 304 Alepidea, 92 Angolluma, 349 Apodicarpum, 103 Aletes, 98 Angoseseli, 100 Apophragma, 472 Aliopsis, 488 Anidrum Apoxyanthera, 318 Allamanda, 277 Anidrum sect.
  • Patterns of Diversity of Floral Symmetry in Angiosperms: a Case Study of the Order Apiales

    Patterns of Diversity of Floral Symmetry in Angiosperms: a Case Study of the Order Apiales

    S S symmetry Article Patterns of Diversity of Floral Symmetry in Angiosperms: A Case Study of the Order Apiales Maxim S. Nuraliev 1,2,* , Dmitry D. Sokoloff 1, Polina V. Karpunina 1 and Alexei A. Oskolski 3,4 1 Faculty of Biology, Lomonosov Moscow State University, 119234 Moscow, Russia; [email protected] (D.D.S.); [email protected] (P.V.K.) 2 Joint Russian-Vietnamese Tropical Scientific and Technological Center, Hanoi 10000, Vietnam 3 Department of Botany and Plant Biotechnology, University of Johannesburg, P.O. Box 524, Auckland Park 2006, Johannesburg, South Africa; [email protected] 4 Botanical Museum, V.L. Komarov Botanical Institute, 197376 St. Petersburg, Russia * Correspondence: [email protected]; Tel.: +7-495-9391827 Received: 16 March 2019; Accepted: 30 March 2019; Published: 3 April 2019 Abstract: Floral symmetry is widely known as one of the most important structural traits of reproductive organs in angiosperms. It is tightly related to the shape and arrangement of floral parts, and at the same time, it plays a key role in general appearance (visual gestalt) of a flower, which is especially important for the interactions of zoophilous flowers with their pollinators. The traditional classification of floral symmetry divides nearly all the diversity of angiosperm flowers into actinomorphic and zygomorphic ones. Within this system, which is useful for ecological studies, many variations of symmetry appear to be disregarded. At the same time, the diversity of floral symmetry is underpinned not only by ecological factors, but also by morphogenetic mechanisms and constraints. Sometimes it is not an easy task to uncover the adaptive or developmental significance of a change of the floral symmetry in a particular lineage.