TELOPEA Publication Date: 27 September 1991 Til
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Volume 4(3): 497–507 TELOPEA Publication Date: 27 September 1991 Til. Ro)'al BOTANIC GARDENS dx.doi.org/10.7751/telopea19914946 Journal of Plant Systematics 6 DOPII(liPi Tmst plantnet.rbgsyd.nsw.gov.au/Telopea • escholarship.usyd.edu.au/journals/index.php/TEL· ISSN 0312-9764 (Print) • ISSN 2200-4025 (Online) 497 Alloxylon (Proteaceae), a new genus from New Guinea and eastern Australia Peter H. Weston and Michael D. Crisp Abstract Weston, Peter H.I, and Crisp, Michael D.2 (1 National Herbarium of New South Wales, Royal Botanic Gardens, Sydney NSW Australia 2000; 2 Australian National Botanic Gardens, GPO Box 1777, Canberra ACT Australia 2601; present address: Division of Botany and Zoology, Australian National University, GPO Box 4, Canberra ACT 2601) 1991. Alloxylon (Proteaceae), a new genus from New Guinea and eastern Australia. Telopea 4(3): 497-507. Oreocallis sens. lat. consists of two distinct clades, one in South America, the other in Australasia, that together are likely to be paraphylet ic. Newly sampled characters strongly support the monophyly of the Australasian group. We describe the new genus Alloxylon to accommodate the Australasian species of Oreocallis sens. lat. and revise its species. Alloxylon fIammeum is described as new and new combinations are made for A. brachycarpum, A. wickhamii and A. pinnatum. Introduction Cladistic analyses of the subtribe Embothriinae of the family Proteaceae (Weston & Crisp 1987, in prep.) show the genus Oreocallis R. Br. sens. lat. to comprise two branches of an unresolved trichotomy. The third branch is the genus Telopea, a well corroborated clade of five species. The sister group to this trichotomy is Embothrium, the only other genus in the Embothriinae. According to these results, Oreocallis is likely to be paraphyletic, necessitating a nomenclatural rearrangement in order to ensure that all formal names apply to monophyletic groups. Two obvious nomenclat ural alternatives are to (a) sink Telopea and Oreocallis into a single genus, or (b) elevate the two distinct groups within Oreocallis sens. lat. to generic rank. We have chosen the latter alternative for the following reasons. Firstly, Telopea and both clades in Oreocallis sens. lat. are more thoroughly characterised by synapomorphies than the group that includes all three of these clades. Option (b) is thus more likely to be nomenclaturally stable than option (a). Secondly, option (b) requires fewer nomen clatural changes than does option (a): only four species are moved into a new genus whereas option (a) requires either that five species of Telopea be moved to Oreocallis or that six species of Oreocallis be moved to Telopea (Telopea and Oreocallis were both established by Brown (1810) and so are equal in priority). Therefore we here establish the new genus Alloxylon to accomodate the three Australian and one New Guinean species formerly included in Oreocallis. Oreocallis sens. strict. comprises two species that broadly overlap in distribution in Peru and Ecuador, O. grandiflora and O. mucronata. Since the publication of our first cladistic analysis of the Embothriinae (Weston & Crisp 1987), knowledge of the wood, fruits, leaf anatomy and seedlings in the EmErn bothriinae has improved considerably, providing a number of new characters for cladistic analysis. Moreover, we now realise that there are three species, one previously unnamed, in north-eastern Australia and New Guinea, not just one as we had previ ously believed, nor two as Sleumer (1954) had thought. The data presented here were compiled from herbarium specimens, cultivated plants, field observations and literature. Specimens held by the following herbaria were 498 Te/opea Vol. 4(3): 1991 examined: BRI, CANB, CBG, MEL, MO, NSW, QRS, SYD. All Australian species are in cultivation at the Royal Botanic Gardens Sydney. Terms denoting form are taken from Stearn (1973), particularly his figure 19. Inflo rescence terminology follows that of Johnson & Briggs (1975) and Weston & Crisp (1987), while leaf venation is described using the terms of Hickey (1973). Flower orientation with respect to the unit inflorescence axis is described as diagonal (opposed to anteroposterior) following Johnson & Briggs (1963, 1975) and Weston & Crisp (1987). Taxonomy Alloxylon P. Weston & Crisp, gen. nov. Arbores parvae ad elatae, in silvis humidis clausis crescentes; cotyledonibus succulentis; bracteis involucralibus nullis; floribus zygomorphis, in alabastro leviter incurvis, roseis ad rubris; perianthio in alabastro plus minusve cylindrico, tubo proximali lineari et limbo distali introrso-obliquo subgloboso, tepalis cohaerentibus postice in tubo fisso post anthesin; polline triporato; pollinophoro valde introrso-obliquo, plus minusve late elliptico ad basin, late conico lateribus concavis, stigmate parvo circulari termi nato; fructu folliculi, mesocarpio interiore lignoso, non late dehiscenti, post dehis centiam cymbiformi; seminibus 6-14, ala seminis basalis longitudine latitudineque loculum fructus fere aequanti sed in seminibus distalibus successive breviore. TYPE: Alloxylon flammeum P. Weston & Crisp. Cotyledons fleshy, epigeal, shortly auriculate, green after germination. First two seedling leaves opposite; all later leaves alternate, with spiral phyllotaxis. Small to tall rainforest trees. Trichomes unbranched. Glandular hairs absent. Adult leaves simple and entire or pinnate with entire leaflets, tapered to petiole. Intermediate leaves simple and entire or deeply pinnatisect or pinnate. Seedling leaves simple, deeply lobed or entire. Conflorescences axillary or terminal, pseudoracemose, later ally bearing flower-pairs (unit inflorescences sensu Johnson & Briggs 1975), basitonic or acrotonic, solitary or aggregated into superconflorescences; involucral bracts absent; unit inflorescences lacking floral bracts and peduncles. Flowers zygomorphic, diag onally oriented with respect to unit inflorescence axis, gently incurved in bud, her maphrodite, ornithophilous, pink to red; torus strongly oblique, lowest on posterior side of flower; perianth ± cylindrical in bud, consisting of proximal linear claw and distal inward-oblique, subglobose limb; segments cohering posteriorly in split tube after anthesis; staminal filaments adnate to perianth segments; anthers circular, introrse, with connective projecting slightly beyond tips of loculi; pollen triporate; hypogynous glands connate, forming posterior, lunate nectary; gynoecium glabrous, c. as long as perianth; gynophore well-developed, half as long as style or more; ovary slightly swollen, containing numerous, laterally attached, winged, anatropous ovules; pollen presenter strongly inward-oblique, ± broad-elliptical at base, broadly conical with concave sides, terminated by small circular stigma. Fruit a follicle, narrow-ellipsoid to narrow-obovoid, with woody inner mesocarp, not opening widely, canoe-like on dehiscence. Seeds 6-14, interleaved by papery, dark brown interseminallayer; em bryo flattened, fleshy; wing distal to embryo, narrow-oblong, truncate to rounded or acute; wing of basal seed nearly as long and wide as loculus of fruit but progressively shorter in distal seeds; raphe entering seed on outside edge of embryo, running along edge onto wing then running diagonally across wing to inside edge near distal tip, sharply reversing and running along inside margin of wing to embryo. Weston & Crisp, Alloxylon 499 Three species endemic in eastern Australia, and one endemic in New Guinea and the Aru Islands. This genus was formerly included in Oreocallis, which, in our delimita tion, comprises two species in Peru and Ecuador. All species, but A. flammeum in particular, are showy flowering trees suitable for cultivation in mildly temperate to tropical areas. The generic name is derived from the Gr~ek allo (other) and xylon (wood), referring to the wood anatomy of the genus, which is markedly different from that of other Proteaceae (Chattaway 1948; Papassotiriou, Weston & Wilkins unpublished data). Key to the species 1 Adult leaves mostly pinnate, some simple; leaf(let) apices attenuate; venation of terminal leaflets and simple leaves eucamptodromous; conflorescences 50-140- flowered, solitary; pollen deep crimson ....................................... 1. A. pinnatum 1* Adult leaves simple and entire; leaf apices obtuse or acute; venation brochidodr omous; conflorescences 4-52-flowered, often grouped in superconflorescences; pollen yellow. 2 Flowers glabrous, dull pinkish red; conflorescences 4- 20-flowered, symmetri- cal to slightly asymmetrical; leaves coriaceous ................... 2. A. wickhamii 2* Flowers covered in (sometimes very short) ferruginous hairs, pinkish red to bright orange-red; conflorescences (2-)10-52-flowered, strongly asymmetrical, the pedicels strongly curved so that all flowers are similarly oriented on the same side of conflorescence axis; leaves chartaceous. 3 Hairs mostly spreading, straight to curly, 0.1-0.5 mm long; young branch- lets densely hairy .......................................... ........................ 3. A. flammeum 3* Hairs appressed, straight, to 0.1 mm long; young branchlets sparsely to moderately hairy ............................................................ 4. A. brachycarpum 1. Alloxylon pinnatum (Maiden & Betche) P. Weston & Crisp, comb. nov. BASIONYM: Embothrium wickhamii W. Hill & F. Muell. var. pinnatum Maiden & Betche, Proc. Linn. Soc. New South Wales 35: 795 (1910). TYPE CITATION: 'Dorrigo (J.L. Boorman; December, 1909).' LECTOTYPE (here designated): NEW