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Archontophoenix (Arecaceae) Description of Species and Notes Article and photos by John Leslie Dowe Australian Center for Tropical Freshwater Research James Cook University. Townsville, QLD Australia [email protected] O f the nine endemic genera of palms that occur in Australia, Archontophoenix is the most diverse with six species. The genus is mainly confined to high rainfall areas, most often in well-developed rainforest, and is intolerant of fire or extended drought. Although predominantly tropical, one species extends to warm temperate Australia. Some of the tropical species occur at relatively high altitudes, in cool wet conditions, and may represent evidence of relictual distribution patterns. Fossil evidence suggests that Arecoid palms were once widespread across southern Australia, but whether they can be directly related to extant genera such as Archontophoenix cannot be verified at present. The morphological variation within Archontophoenix is relative narrow, and species are primarily delimited by the following characters: - leaflets with the presence/absence of dense silvergrey tomentum, and ramenta on the abaxial surface - stamen number - fruit size and colour - condition of the fibres in the mesocarp - colouration and morphology of the crownshaft The following treatment is based on extensive field-work and herbarium research conducted by the author, and is largely taken from the forthcoming treatment of the palm family in the Flora of Australia series and the ‘Systematics of the Australian Palm Flora’ project. Taxonomy and species descriptions Archontophoenix H.Wendl. & Drude, Linnaea 39: 182, 211 (1875); from the Greek archon (king or ruler), and phoenix (palm). Loroma O.F.Cook, J. Wash. Acad. Sci. 5: 117 (1915). Solitary, erect palms. Trunk slender, frequently with swollen base. Leaves paripinnate, caducous; rachis ±straight or curved; leafsheath forming a tubular crownshaft. Leaflets reduplicate, linear-acute, in a single plane; midrib prominent; abaxial surface green or with silver-grey scales; ramenta present or lacking. Inflorescence infrafoliar, a divaricate panicle branched to 3 or 4 orders; bracts 2, fully enclosing inflorescence in bud; prophyll fibrous; peduncular bract membranous, inserted slightly above attachment of prophyll; rachis longer than peduncle; rachillae flexuose. Flowers in triads Figure 1 (left) - Archontophoenix alexandrae, Lamb Range, north-east Queensland. species leaflet abaxial (lower) surface fruit shape and length mesocarp fibres A. alexandrae silver–grey scales, ramenta lacking ovoid–globose, 8–14 mm mesocarp fibres to 1 mm wide, flat, sparingly branched, remaining tight in the dried state A. cunninghamiana no silver–grey scales, ramenta present ovoid–globose, 10–15 mm mesocarp fibres to 1 mm wide, flat, branched, loosening in the dried state A. maxima silver–grey scales, ramenta lacking conic–ovoid, 11–15 mm mesocarp fibres to 1 mm wide, flat, sparingly branched A. myolensis silver–grey scales, ramenta lacking conic–ovoid– elongate, 13–21 mm mesocarp fibres terete or flat, thin to thick, to 0.5 mm wide, branched, forked about midway A. purpurea silver–grey scales, ramenta present ellipsoid to globose, 20–26 mm fibres flat, to 2 mm wide, branched, overlaid and interspersed with thin fibres A. tuckeri silver–grey scales, ramenta lacking ovoid, 15–25 mm fibres in two layers, the outer layer of fibres thin, occasionally branched, the inner layer thick, flat, to 3 mm wide, mostly branched; fibres compact in the dried state Table 1. Distinguishing characters for species of Archontophoenix. Characters unique to a single species are highlighted in bold italics. throughout or with paired or solitary male flowers distally on the rachillae, spirally arranged. Staminate flower asymmetric in bud; stamens 9–35; anthers dorsifixed; filaments curved or deflexed. Pistillate flower smaller than staminate. Fruit moderate, conic-ovoid to subglobose, red; stigmatic remains apical to sub-apical; epicarp thin, smooth; mesocarp with anastomosing fibres. Seed ovoid to subglobose; endosperm ruminate; embryo basal. Eophyll bifid. Archontophoenix is a genus of six species endemic to Australia. Three of the species are narrow endemics that are confined to either a single drainage system (A. myolensis), or mountain ranges or massifs (A. maximus, A. purpurea), while the other three have widespread distribution, one confined to Cape York Peninsula (A. tuckeri), one to eastern Queensland (A. alexandrae) and the other (A. cunninghamiana) to central Queensland south to southern New South Wales. Species occur primarily as canopy emergents or subcanopy elements in closed forests in high rainfall areas on various soil types at low to moderate elevation. Archontophoenix alexandrae and A. cunninghamiana co–occur in the Eungella Range, where both grow at high elevation in intermingled populations. There has been no recorded evidence of hybridity between these species in these natural populations, although hydridity may be achieved by controlled pollination in cultivation. Characters that distinguish species are presented in Table 1. Archontophoenix alexandrae (F.Muell.) H.Wendl. & Drude, Linnaea 39: 212 (1875) Synonyms: Ptychosperma alexandrae F.Muell. Archontophoenix alexandrae var. schizanthera H.Wendl. & Drude Archontophoenix veitchii H.Wendl. & Drude Ptychosperma beatriceae F.Muell. Archontophoenix beatriceae (F.Muell.) F.M.Bailey Archontophoenix alexandrae var. beatriceae (F.Muell.) C.T.White ex L.H.Bailey Jessenia glazioviana Dammer Trunk to 30 m tall, to 30 cm diam. Leaves 10–12, to 4.5 m long, becoming vertically orientated; leafsheath mid to lime green. Leaflets 60-80 each side of rachis, to 75 cm long, dark green adaxially, silver-grey abaxially; ramenta lacking. Inflorescence to 100 cm long, branched to 4 orders; axes The Palm Journal #192 5 Key to species Perianth lilac to purple–mauve; leaflets dark green on both surfaces . . . . . . . . . . . . . . . . . . . . . A. cunninghamiana Perianth white to cream; leaflets dark green adaxially, silver-grey abaxially Mesocarp fibres less than 0.5 mm wide, unravelling in the dried state . . . . . . . . . . . . . . . . . . . A. myolensis Mesocarp fibres to greater than 0.5 mm wide, remaining compact in the dried state Leaflets with ramenta on abaxial midrib . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. purpurea Leaflets lacking ramenta on abaxial midrib Flowers closely spaced on rachillae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. tuckeri Flowers widely spaced on rachillae Floral pits with margins round and low; raphe fibres in fruit strongly adherent . . . . A. maxima Floral pits with raised sharp margins; raphe fibres in fruit weakly adherent . . . . . A. alexandrae white to creamy green; rachillae to 30 cm long, erect to semi-pendulous, moderately flexuose distally. Flowers widely spaced on rachilla; floral pits with raised sharp margins; perianth white to cream; staminate flower to 9.5 mm long; stamens 9–16; filaments curved; pistillode about equal to petals; pistillate flower to 4 mm high. Fruit ovoid, 8–14 mm long, 6–11 mm wide; stigmatic remains apical; mesocarp fibres to 1 mm wide, flat, sparingly branched, remaining compact in the dried state. Seed globose, 8 mm diam.; raphe fibres weakly adherent. Figure 1. Common names: Alexandra Palm, King Palm. Occurs in coastal Queensland from Melville Range south to Baffle Ck. Grows on various soil types in rainforest, swamp forest and moist sclerophyll forest at 0–1000 m alt. Flowers all year; fruits all year. Named for Princess Alexandra of Denmark (1844–1925), daughter of King Christian IX and Louise of Hessen–Kassel, married to King Edward VII of Great Britain (1841–1910) in 1863. Archontophoenix alexandrae is the most widespread and prevalent palm in north-eastern Australia, contributing to a characteristic landscape physiognomy, although most of its former lowland habitat has been cleared for agriculture. Large populations can be seen in the ‘Wet Tropics Bioregion’, and it is well conserved in national parks and conservation reserves. It most closely resembles A. maxima, from which it differs by a less robust inflorescence and larger fruits, and A. myolensis, which is distinguished by elongate fruit with narrow, rather than wide, fibres in the mesocarp. These three species have the abaxial surface of the leaflets with dense silver–grey scales, and lack ramenta. 6 The Palm Journal #192 Archontophoenix cunninghamiana (H.Wendl.) H.Wendl. & Drude, Linnaea 39: 214 (1875) Synonyms: Ptychosperma cunninghamianum H.Wendl. Seaforthia cunninghamii (H.Wendl.) Hort. ex F.M.Bailey Loroma cunninghamiana (H.Wendl.) O.F.Cook Jessenia amazonum Drude Loroma amethystina O.F.Cook Seaforthia nobilis Lhotsky Trunk to 30 m tall, to 30 cm diam. Leaves 9–12, to 4.5 m long, becoming vertically orientated; leafsheath dark green to green-purple. Leaflets 70-90 each side of rachis, to 100 cm long, dark green on both surfaces; ramenta present. Inflorescence to 150 cm long, branched to 4 orders; axes white to yellow green becoming mauve in infructescence; rachillae to 55 cm long, pendulous, moderately flexuose distally. Flowers closely spaced on rachilla; floral pits with raised sharp margins; perianth lilac to purple–mauve; staminate flower to 6 mm long; stamens 9–18; filaments deflexed; pistillode about equal to petals; pistillate flower to 4 mm high. Fruit ovoid, 10–15 mm long, 6–11 mm wide; stigmatic remains apical; mesocarp fibres to 1 mm wide, flat, sparingly branched, remaining compact in the dried state. Seed globose, to 9 mm diam.; raphe fibres embedded, strongly adherent. Figure 2. Common names: Bangalow Palm, Piccabeen Palm. Endemic to coastal Queensland and New South Wales, from Mt Elliot (south of Townsville) to Durras Mountain. Grows on various soil types in rainforest at 0–1200 m alt. Flowers all year; fruits all year. Named for Allan Cunningham (1791–1839), British Figure 2 - Archontophoenix cunninghamiana, Kondalilla Falls, south-east Queensland. globose, 6 mm diam.; raphe fibres deeply embedded and strongly adherent. Figure 3. Common names: Walsh River Archontophoenix, Herberton Range Archontophoenix. Endemic to the Herberton Range and Mt Haig Range, western Atherton Tablelands, north–east Queensland. Grows on basaltic soils in rainforest and moist sclerophyll forest at 800–1200 m alt. Flowers June–Aug,; fruits Dec.–Mar. Named in reference to the larger size of the inflorescence, and in particular the greater width of the peduncle as compared to other species of Archontophoenix. Archontophoenix maxima is distinguished by the robust peduncle and the leaves which are held only moderately laterally twisted, and not strongly so as in other species of Archontophoenix. It most closely resembles A. alexandrae. Archontophoenix myolensis Dowe, in J.L.Dowe & D.R.Hodel, Austrobaileya 4(2): 237, fig 1B (1994) Figure 3 - Archontophoenix maxima, north-east Queensland. botanist, who collected extensively in Australia, arriving in 1816 as ‘King’s Botanist to the Royal Botanic Gardens Kew’. The relationships of Archontophoenix cunninghamiana to others in the genus are not clear; it is otherwise characterized by leaflets that lack silver–grey scales, and the presence of ramenta on the mid of the abaxial surface of the leaflets. Sepals and petals of both staminate and pistillate flowers are uniquely lilac to purple–mauve. Archontophoenix maxima Dowe, in J.L.Dowe & D.R.Hodel, Austrobaileya 4(2): 235, fig 1A (1994) Trunk to 22 m tall, to 30 cm diam. Leaves 8–11, to 5 m long, becoming obliquely orientated; leafsheath mid green. Leaflets c. 85 each side of rachis, to 90 cm long, dark green adaxially, silver-grey abaxially; ramenta lacking. Inflorescence to 140 cm long, branched to 4 orders; axes cream-green; rachillae to 40 cm long, erect to semipendulous, moderately flexuose distally. Flowers widely spaced on rachilla; floral pits with rounded low margins; perianth white to cream; staminate flower to 6 mm long; stamens 11–16; filaments curved; pistillode about equal to petals; pistillate flower to 5 mm high. Fruit subglobose to ovoid, 11–15 mm long, 7–12 mm wide; stigmatic remains apical; mesocarp fibres to 1 mm wide, flat, sparingly branched, remaining compact in the dried state. Seed 8 The Palm Journal #192 Trunk to 20 m tall, to 30 cm diam. Leaves 9–12, to 4 m long, becoming vertically orientated; leafsheath blue green. Leaflets 68-71 each side of rachis, to 110 cm long, dark green adaxially, silver-grey abaxially; ramenta lacking. Inflorescence to 160 cm long, branched to 4 orders; axes white to creamy green; rachillae to 30 cm long, erect to semi-pendulous, moderately flexuose distally. Flowers closely spaced on rachilla; floral pits with margins not prominant; perianth white to cream; staminate flower to 9 mm long; stamens 14–24; filaments curved; pistillode about equal to petals; pistillate flower to 5 mm high. Fruit elongate, conicovoid to ovoid, 13–21 mm long, 10–12 mm wide; stigmatic remains apical to slightly subapical; mesocarp fibres terete, to 0.5 mm wide, sparingly branched, becoming unravelled in the dried state. Seed ovoid-elongate, 14–20 mm long, 8–9.5 mm diam., raphe fibres strongly adherent. Figure 4. Common name: Myola Archontophoenix. Endemic to the Myola district, eastern Atherton Tablelands, north-east Queensland. Grows on soils derived from metamorphic rocks in rainforest at 350–400 m alt. Flowers May–July; fruits Dec.–Mar. Named for the locality of Myola, which is located about 5 km west of the township of Kuranda, Atherton Tableland, Queensland. Archontophoenix myolensis is the rarest species of Archontophoenix, confined to the Warril Creek drainage system and nearby slopes, at 350–400 m elevation, on the Atherton Tableland, west of Kuranda township, Queensland. The population consists of only a few hundred mature individuals, of which only a small percentage is conserved in National Parks or nature reserves. The species’ rarity and its population being within an area of urban expansion, places it under considerable threat and it has an Endangered conservation status. It has been reported that the fruit and seedling Figure 4 Archontopehoenix myolensis, Warril Creek, Myola, north-east Queensland. The Palm Journal #192 9 morphologies are adaptations to riparian conditions: the fibres in the mesocarp disassociate in the dried state and act as small hooks, which enables the seeds to harness themselves amongst debris or within sediments, and the seedling leaves become pinnate at a much earlier stage than do other species of Archontophoenix [which suggests a possible adaptation to fast flowing water]. The species is distinguished by the elongate conic-ovoid to ovoid fruit, and with narrow, rather than wide, fibres in the mesocarp. Archontophoenix purpurea Hodel & Dowe, in J.L.Dowe & D.R.Hodel, Austrobaileya 4(2): 238, fig. 1C (1994) Trunk to 25 m tall, to 45 cm diam. Leaves 7–15, to 6 m long, becoming obliquely orientated; leafsheath dark green to purple. Leaflets 5585 each side of rachis, to 105 cm long, dark green adaxially, silver-grey abaxially; ramenta present on abaxial midrib. Inflorescence to 135 cm long, branched to 4 orders; axes green; rachillae to 85 cm long, erect to semipendulous, irregularly flexuose distally. Flowers closely spaced on rachilla; floral pits with raised sharp margins; perianth white to cream; staminate flower to 7 mm long; stamens 20–35; filaments curved; pistillode about equal in length to petals; pistillate flower to 6.5 mm high. Fruit ovoid to globose, 20–26 mm long, 8–22 mm wide, red-crimson at maturity; stigmatic remains apical; mesocarp fibres to 2 mm wide, flat, sparingly branched, interspersed with terete thin fibres, remaining compact in the dried state. Seed subglobose, to 23 mm long, to 18 mm diam.; raphe fibres strongly adherent. Figure 5. Common name: Mount Lewis Archontophoenix. Endemic to north-east Queensland, on Mts Finnigan, Spurgeon and Lewis. Grows on granitic soils in rainforest at 400–1200 m alt. Flowers Oct.–Apr.; fruits Sept.–Apr. Named for the distinctive purple colouration of the crownshaft - Latin purpureus. Archontophoenix purpurea is one of the most distinctive species in the genus, having a prominent purple crownshaft and large fruit. The colouration of the crownshaft is due to three separate components of surface indumentum. The innermost layer consists of the dark blue–grey surface of the crownshaft, a powdery glaucous bloom, and a dense arrangement of reddish mahogany coloured scales. From a distance the three components merge to form the characteristic purple colour. The fruit, at 20–26 mm long, are the largest in the genus. They are red–crimson at maturity, have prominent apical stigmatic remains, and the mesocarp fibres are up to 2 mm wide. In addition, the leaflets have a unique combination of possessing both silver–grey scales and ramenta on the abaxial surface. Archontophoenix tuckeri Dowe, in J.L.Dowe & D.R.Hodel, Austrobaileya 4(2): 240, fig. 1D (1994) Figure 5 - Archontophoenix purpurea, Mt Lewis, ca. 1100 m altitude, north-east Queensland. Trunk to 20 m tall, to 26 cm diam. Leaves 7–12, to 3.15 m long, held flat or becoming moderately obliquely orientated; leafsheath lime green. Leaflets 43-68 each side of rachis, to 93 cm long, dark green adaxially, silver-grey abaxially; ramenta lacking. Inflorescence to 60 cm long, branched to 3 orders, axes green; rachillae to 30 cm long, erect to semi-pendulous, strongly flexuose throughout. Flowers closely spaced on rachilla; floral pits with raised sharp margins; perianth white; staminate flower to 7 mm long; stamens 13–19; filaments curved; pistillode about equal in length to petals; pistillate flower to 5 mm high. Fruit ovoid, 15–25 mm long, 10–15 mm wide; stigmatic remains apical; mesocarp fibres two-layered, outer layer thin, terete, occasionally branched, inner layer to 3 mm wide, flat, branched, remaining compact in the dried state. Seed globose, 9 mm diam.; raphe fibres weakly adherent. Figure 6. Common names: Rocky River Archontophoenix, Peach Creek Archontophoenix, Iron Range Archontophoenix. Endemic to north-east Queensland, on Cape York Peninsula, Lockerbie Scrub, Iron Range and McIlwraith Range. Grows on various soil types in rainforest at 0–500 m alt. Flowers June–Feb.; fruits Aug.–Nov. Named for Robert James Thomas Tucker (b.1955–d.1992), horticulturist, botanical illustrator, taxonomist (Pandanaceae) and landscape designer for Townsville Botanic Gardens, 1982–1992, and designer of The Palmetum, Townsville, a botanic garden devoted to palms. Tucker collected palms and pandanus throughout Cape York Peninsula and was the author of The Palms of Subequatorial Queensland. Archontophoenix tuckeri is distinguished by the narrow elongate light green crownshaft and large bright red fruit. Otherwise, habit and general appearance is similar to A. alexandrae. The fruit, 15–25 mm long, have been observed to have an epicarp that ‘cracks’ at maturity, thus exposing the mesocarp. Figure 6 - Archontophopenix tuckeri, West Claudie River, Iron Range, north Queensland. The Palm Journal #192 11