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Inclusion of the endemic New Caledonian genus Pseudosciadium in Delarbrea (Apiales, Myodocarpaceae) Porter P. LOWRY II Missouri Botanical Garden, P.O. Box 299, St. Louis, MO 63166-0299 (USA) [email protected] and Département Systématique et Évolution (USM 602), Muséum national d’Histoire naturelle, CP 39, 57 rue Cuvier, F-75231 Paris cedex 05 (France) [email protected] Gregory M. PLUNKETT Virginia Commonwealth University, P.O. Box 482012 Richmond, VA 23284-2012 (USA) [email protected] Virginie RAQUET Département Systématique et Évolution (USM 602), Muséum national d’Histoire naturelle, CP 39, 57 rue Cuvier, F-75231 Paris cedex 05 (France) [email protected] Taylor S. SPRENKLE Virginia Commonwealth University, P.O. Box 482012 Richmond, VA 23284-2012 (USA) Joël JÉRÉMIE Départements Milieux et Peuplements aquatiques (USM 403) et Systématique et Évolution (USM 602), Muséum national d’Histoire naturelle, CP 39, 57 rue Cuvier, F-75231 Paris cedex 05 (France) [email protected] ABSTRACT Recent molecular and morphological phylogenetic analyses of the newly- recognized family Myodocarpaceae (Apiales) have shown that the genus Pseudosciadium (represented by a single species endemic to New Caledonia) is nested within a well-supported clade that also comprises members of the KEY WORDS Apiales, genus Delarbrea, necessitating its inclusion within a more broadly circumscri- Myodocarpaceae, bed Delarbrea. An expanded description is provided for Delarbrea along with Pseudosciadium, a detailed description of the newly transferred species, D. balansae, accompa- Delarbrea, New Caledonia, nied by full citation of specimens, an illustration and a preliminary conserva- conservation. tion assessment calculated according to IUCN Red List criteria. ADANSONIA, sér. 3 • 2004 • 26 (2) : 251-256 © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. 251 Lowry II P.P., Plunkett G.M., Raquet V., Sprenkle T.S. & Jérémie J. RÉSUMÉ Inclusion du genre néocalédonien Pseudosciadium dans Delarbrea (Apiales, Myodocarpaceae). Des analyses phylogénétiques moléculaires et morphologiques de la famille des Myodocarpaceae (Apiales), reconnue récemment, ont montré que le genre Pseudosciadium (représenté par une unique espèce endémique de Nouvelle- Calédonie) est inclu à l’intérieur d’un clade bien soutenu qui renferme aussi des membres du genre Delarbrea, ce qui nécessite son placement au sein de ce MOTS CLÉS Apiales, dernier genre plus largement délimité. Une description élargie est donnée Myodocarpaceae, pour Delarbrea ainsi qu’une description détaillée de l’espèce D. balansae, Pseudosciadium, accompagnée de la liste du matériel étudié, d’une illustration et d’une analyse Delarbrea, Nouvelle-Calédonie, préliminaire du statut de conservation selon les critères des Listes Rouges de conservation. l’UICN. The relationships and placement of the mono- rowed at the base, where it joined two other gen- typic, endemic New Caledonian genus Pseudo- era, Apiopetalum Baill. from New Caledonia and sciadium Baill. have confounded botanists for Mackinlaya F. Muell. from Australia and New more than 125 years. BAILLON (1878, 1879) orig- Guinea. The affinities of Pseudosciadium remained inally described the genus on the basis of a beau- ambiguous, however, until mature fruits became tifully preserved flowering collection made by available demonstrating unmistakable homolo- BALANSA in 1871 at the mouth of the Dothio gies with the fruits of Delarbrea and Myodocarpus, River just north of the town of Thio along the including the presence of distinctive oil vesicles in eastern coast of New Caledonia. At the time, the endocarp, a feature known nowhere else in BAILLON indicated that Pseudosciadium was the family. This prompted LOWRY (1986a) to closely related to (and perhaps intermediate suggest that Delarbrea, Myodocarpus and between) two other genera then placed in Pseudosciadium form a monophyletic group Araliaceae, Delarbrea Vieill. and Myodocarpus within Araliaceae, signaling a return to BAILLON’s Brongn. & Gris, both of which were regarded as (1878, 1879) original concept. endemic to New Caledonia. BAILLON’s initial Over the years the position of these three gen- intention had been to describe this new taxon as a era within Apiales has likewise presented diffi- species of Myodocarpus, but the presence in some culties. Because Delarbrea, Myodocarpus and flowers of valvate petals with a narrowed base Pseudosciadium are all woody, they have tradition- lacking an evident claw (vs imbricate, distinctly ally been placed in Araliaceae, despite the presence clawed petals in Myodocarpus) and of an evident of several features generally regarded as more typi- stipe at the base of the ovary (lacking in cal of Apiaceae, including a bicarpellate gynoe- Myodocarpus) prompted him to recognize it as a cium, petals with inflexed tips and, in the case of distinct genus. Subsequent authors (e.g., HARMS Myodocarpus, a schizocarpic fruit. To clarify the 1894-1897; CALESTANI 1905; VIGUIER 1906, affinities of these and several other taxa often 1925; HUTCHINSON 1967), placing substantial regarded as “intermediates” between the two fami- emphasis on corolla aestivation for the delimita- lies, PLUNKETT & LOWRY (2001) used a combina- tion of major groups within Araliaceae, separated tion of nuclear (ITS) and plastid (matK) sequence Pseudosciadium from Delarbrea and Myodocarpus data to evaluate the phylogenetic relationships of (often regarded as comprising tribe Myodo- these problematic taxa. Their results confirmed carpeae), and transferred it to tribe Mackinlayeae, that Delarbrea, Myodocarpus and Pseudosciadium defined to include taxa with valvate petals nar- comprise a strongly supported monophyletic 252 ADANSONIA, sér. 3 • 2004 • 26 (2) Inclusion of Pseudosciadium in Delarbrea (Apiales: Myodocarpaceae) group segregated from the core members of Porospermum F. Muell., Fragm. 7: 94 (1870). — Araliaceae and from other basally branching lin- Type: Porospermum michieanum F. Muell. Pseudosciadium Baill., Adansonia 12: 130 (1878), syn. eages in Apiales, a conclusion that led DOWELD nov. — Type: Pseudosciadium balansae Baill. (2001) to describe Myodocarpaceae as a distinct family. Within this clade, two well supported sub- Monocaulous or sparsely branched, glabrous, clades were identified, one comprising Delarbrea + unarmed treelets or small trees. Stem tips, peti- Pseudosciadium (which share terete fleshy fruits oles and base of inflorescence often glaucescent. and noncalyptrate corollas) and another contain- Leaves imparipinnate, alternate, clustered at ends ing species of Myodocarpus (characterized by dor- of branches; leaflets opposite to subopposite (or sally flattened, winged schizocarpic fruits and sometimes alternate, especially the lower ones), buds with calyptrate corollas) a finding supported entire or remotely dentate, occasionally serrate on by data from wood anatomy (OSKOLSKI et al. the first few leaves borne on new shoots (to 1997). The study of PLUNKETT & LOWRY (2001) deeply serrate or lacerate in juvenile foliage of was not, however, able to resolve the relationship some species), the base of the lateral leaflets between Delarbrea and Pseudosciadium, although oblique; rachis not articulated; petiole with an some of their cladograms suggested that expanded, clasping base with membranous or Pseudosciadium may be nested within the scarious margins. Inflorescence a panicle of um- Delarbrea subclade. In a subsequent study, bellules, terminal, erect or pendant, the lateral SPRENKLE (2002; see also PLUNKETT et al. 2004) branches, peduncles and umbellules subtended carried out a more detailed phylogenetic analysis by membranous, scarious or foliaceous bractlets, focusing specifically on relationships within the pedicels free or basally united into groups of Myodocarpaceae using an expanded set of 2-4, articulated below the ovary. Flowers her- molecular data from three nuclear markers (ITS, maphrodite and protandrous, often also func- ETS and the 5S non-transcribed spacer) and one tionally staminate (in andromonoecious species), chloroplast marker (trnL-trnF). The results of that actinomorphic. Sepals united below into a short study confirmed the earlier finding of two well- tube, the 5 free lobes obtuse or triangular to differentiated clades within the family, and clearly rounded or oblate, valvate, sometimes with a showed that Pseudosciadium falls within the scarious margin, not or only slightly expanding in Delarbrea clade. In a complementary phylogenetic fruit, the margins often somewhat scarious. Petals study using 35 morphological characters, RAQUET 5, imbricate or valvate, broadly ovate to obovate (2004) also confirmed the presence of two distinct or spatulate, keeled within, narrowed to distinctly lineages within Myodocarpaceae corresponding to clawed toward the base. Stamens 5, inflexed Myodocarpus and Delarbrea + Pseudosciadium, as before anthesis, the filaments stout, the anthers well as the placement of Pseudosciadium nested with 4 thecae, cream-white, dorsifixed. Ovary among the members of Delarbrea. The morpho- inferior, 2-carpellate, vestigial in staminate flow- logical analysis also revealed the presence of a stipe ers, sometimes prolonged below into a slender at the base of the fruits of several species of stipe, surmounted by a small, rounded to Delarbrea, further supporting a close relationship depressed conic nectar disc, styles 2, free, erect with Pseudosciadium. Taken together, these find- and appressed at anthesis, spreading as the ± ings indicate that Pseudosciadium
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  • Systematic Wood Anatomy of Myodocarpus, Delarbrea, and Pseudosciadium (Araliaceae)

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

    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.