COMMELINIDAE Takht.

Takhtajan, Sist. Filog. Cvetk. Rast. [Syst. Phylog. Magnolioph.]: 171. 4 Feb 1967


[Arecaceae+Dasypogonaceae+[Commelinanae+Cyperales]]


ARECACEAE Bercht. et J. Presl

Berchtold et Presl, Přir. Rostlin: 266. Jan-Apr 1820, nom. cons.

Palmae Juss., Gen. Plant.: 37. 4 Aug 1789, nom. cons. et nom. alt.; Rhapidaceae Bercht. et J. Presl, Přir. Rostlin: 266. Jan-Apr 1820 [‘Rhapideae’]; Borassaceae Schultz Sch., Nat. Syst. Pflanzenr.: 318. 30 Jan-10 Feb 1832 [’Borasseae’]; Cocosaceae Schultz Sch., Nat. Syst. Pflanzenr.: 316. 30 Jan-10 Feb 1832 [’Cocoineae’]; Coryphaceae Schultz Sch., Nat. Syst. Pflanzenr. 317. 30 Jan-10 Feb 1832; Sabalaceae Schultz Sch., Nat. Syst. Pflanzenr.: 317. 30 Jan-10 Feb 1832 [’Sabalineae’]; Sagoaceae Schultz Sch., Nat. Syst. Pflanzenr.: 316. 30 Jan-10 Feb 1832 [’Sagoineae’]; Phoenicaceae Burnett, Outl. Bot.: 395, 1155. Feb 1835; Phoenicales Burnett, Outl. Bot.: 1155. Feb 1835 [’Phoeniciales’]; Calamaceae Kunth ex Perleb, Clav. Class.: 13. Jan-Mar 1838 [’Calameae’]; Lepidocaryaceae Mart., Hist. Nat. Palm. 3: 196. 23 Sep 1838 [’Lepidocaryinae’]; Phytelephantaceae Mart. ex Perleb, Clav. Class.: 11. Jan-Mar 1838 [’Phytelephanteae’]; Arecales Bromhead in Mag. Nat. Hist., ser. II, 4: 333. Jul 1840; Phoenicopsida Brongn., Enum. Plant. Mus. Paris: xv, 15. 12 Aug 1843 [’Phoenicoideae’]; Nypaceae Brongn. ex Le Maout et Decne., Traité Général Bot.: 624. Jan-Apr 1868 [’Nipaceae’]; Ceroxylaceae Vines, Stud. Text-book Bot. 2: 544. Mar 1895 [’Ceroxylinae’]; Manicariaceae O. F. Cook in Contr. U.S. Natl. Herb. 13: 140. 22 Jun 1910; Acristaceae O. F. Cook in Contr. U.S. Natl. Herb. 16: 252. 14 Mai 1913; Chamaedoreaceae O. F. Cook in Contr. U.S. Natl. Herb. 16: 252. 14 Mai 1913 [’Chamaedoraceae’]; Geonomataceae O. F. Cook in Contr. U.S. Natl. Herb. 16: 252. 14 Mai 1913 [’Geonomaceae’]; Iriarteaceae O. F. Cook et Doyle in Contr. U.S. Natl. Herb. 16: 225. 13 Feb 1913; Malortieaceae O. F. Cook in Contr. U.S. Natl. Herb. 16: 252. 14 Mai 1913; Pseudophoenicaceae O. F. Cook in Contr. U.S. Natl. Herb. 16: 243. 14 Mai 1913; Synechanthaceae O. F. Cook in Contr. U.S. Natl. Herb. 16: 252. 14 Mai 1913; Cocosales Nakai, Hisi-Shokubutsu: 48. 1930 [’Cocoales’]; Arecanae Takht., Sist. Filog. Cvetk. Rast. [Syst. Phylog. Magnolioph.]: 525. 4 Feb 1967; Arecidae Takht., Sist. Filog. Cvetk. Rast. [Syst. Phylog. Magnolioph.]: 525. 4 Feb 1967

Genera/species 180–185/2.425–2.500

Distribution Chiefly pantropical, some species in subtropical regions and a few species in warm-temperate areas.

Fossils The oldest records of Arecaceae leaves and pollen date from the Coniacian and the Santonian. Stems, leaves, seeds, pollen grains (e.g. Echimonocolpites, Longapertites, Mauritiidites, Monocolpites and Spinizonocolpites) and fruits are known from the Maastrichtian onwards, and especially from the Paleocene and Eocene. Flowers of possibly arecaceous origin have been found in Campanian to Maastrichtian layers in Portugal. Fossil records of Nypa and the extinct Nipadites usually consist of pollen grains (Spinizonocolpites) and date back to the Maastrichtian. Numerous palm fossils, such as Arecoideostrobus, Hyphaeneocarpon, Nipadites, Palmocarpon, Palmoxylon, Parapalmocaulon and Sabalophyllum, are known from the Maastrichtian Deccan Intertrappean Beds in India. These consist of stems, leaves, fruits, etc. Finally, a large number of fossils have been identified as belonging in extant genera.

Habit Usually monoecious, polygamomonoecious or dioecious; male and female flowers isomorphic or heteromorphic (sometimes bisexual); evergreen, woody (trees, shrubs or lianas), usually with monopodial growth (rarely branched). Roots or leaflets sometimes modified into spines (spine roots). Aerial roots (stilt roots, prop roots or pneumatophores) present in many species. Nypa consists of mangrove plants. Hyphaene has dichotomous branching.

Vegetative anatomy Phellogen? Wood consisting of primary tissue originating in apical meristem; without or almost without secondary lateral growth. Endodermal cell walls with annular or U-shaped thickenings. Cambium absent (stem sometimes increasing in thickness due to primary growth). Vascular bundles scattered in stem parenchyma. Secondary lateral growth absent (sometimes weak lateral growth by divisions in ground tissue). Vessels present in root, stem and leaves (vessels often very long, up to several metres). Vessel elements with scalariform or simple perforation plates (simple perforation plates absent in, e.g., Nypa); lateral pits? Imperforate tracheary xylem elements tracheids, septate or non-septate; fibres partly associated with xylem and phloem, partly scattered in parenchyma. Wood rays? Axial parenchyma? Wood sometimes fluorescent. Sieve tube plastids usually P2cs type, with cuneate protein crystals and starch grains (rarely P2c type, with cuneate protein crystals, without starch or protein filaments, or P2cfs, with cuneate protein crystals, starch and protein filaments). Nodes multilacunar with several leaf traces. Parenchyma cells often sclerified or lignified (in some groups, e.g. Metroxylon, starchy). Special cells, isodiametric stegmata connected to fibres, with cap-shaped or spherical and often spinulate silica bodies. Calciumoxalate usually as raphides (rarely as crystal sand or solitary crystals).

Trichomes Lepidote and peltate hairs abundant.

Leaves Alternate (rarely distichous or tristichous), seemingly pinnately compound (rarely repeatedly pinnately compound) or seemingly palmately compound (palmate or costapalmate with segment), or simple, pinnately or palmately lobed (rarely entire or bifid), usually differentiated into pseudopetiole and pseudolamina; hastulae often present at junction between pseudopetiole and pseudolamina; rhachis in some species prolonged and modified into whip-like tendril, cirrhus, with plicate, induplicate (splitting along adaxial folds) or reduplicate (splitting along abaxial folds; rarely splitting between folds or non-splitting) ptyxis; leaflets reduplicate: Λ-shaped, or induplicate: V-shaped. Leaves often very large (in Raphia up to more than 25 m). Stipules absent; leaf sheath well developed, closed, often long and wide (sometimes with prickles or spines developed from modified adventitious roots or leaflets), in some species with a ligule-like structure in transition zone to pseudopetiole or pseudolamina. Central pseudopetiole vascular bundles with one or two phloem strands. Venation parallelodromous (seemingly pinnate or palmate). Stomata usually tetracytic; subsidiary cells with oblique divisions. Cuticular wax crystalloids as longitudinally aggregated rodlets (Strelitzia type), chemically dominated by wax esters. Mesophyll with idioblasts containing calciumoxalate as raphides, crystal sand and/or solitary prismatic crystals. Epidermal cells often with silica bodies. Leaf margin usually entire (rarely serrate).

Inflorescence Usually axillary (sometimes terminal), simple or compound panicle, spike- or spadix-like surrounded by one or several spatha-like bracts (inflorescence in some species of Calamus transformed into a climbing flagellum). Partial inflorescences of various shape, usually cymose. Prophyll bicarinate. Floral prophylls (bracteoles) on branches lateral (ultimate inflorescence units cincinni).

Flowers Usually actinomorphic (rarely zygomorphic). Hypogyny. Tepals usually 3+3 (sometimes 2+2; rarely several whorles or up to ten spiral tepals), usually with imbricate (sometimes valvate; inner tepals in, e.g., Cocos with valvate) aestivation, free or more or less connate (in, e.g., Coccothrinax all tepals connate into tube); outer tepals usually sepaloid; inner tepals usually petaloid. Nectar secreted from septal nectaries or androecial nectaries at staminal bases. Disc absent.

Androecium Stamens usually 3+3 (rarely three, antesepalous or antepetalous, sometimes more than six, in Phytelephas and Ammandra to more than 1.000). Filaments usually narrow, erect or inflexed in bud, free or connate at base into tube, free from or more or less adnate to tepals. Anthers basifixed or dorsifixed, usually straight (rarely spirally twisted), often versatile, tetrasporangiate, introrse or latrorse (rarely extrorse), usually longicidal (dehiscing by longitudinal slits; rarely poricidal, dehiscing by apical pores). Tapetum secretory, with binucleate cells. Female flowers usually with few to numerous staminodia, free or connate (sometimes adnate to perianth or gynoecium; sometimes absent).

Pollen grains Microsporogenesis simultaneous (by centripetal or centrifugal cell wall production) or successive. Pollen grains monoporate or diporate, monosulcate, trichotomosulcate, zonosulcate etc. (17 different aperture types found, very rarely inaperturate), shed as monads, bicellular at dispersal. Exine tectate, semitectate or intectate, with columellate to granular infratectum, perforate, reticulate, fossulate or scrobiculate, rugulate, gemmate or with other supratectal processes, or smooth.

Gynoecium Carpels (one to) three or four (to ten), free or more or less connate. Ovary superior, usually trilocular or unilocular (apocarpy) with one fertile locule (pseudomonomerous; rarely up to decemlocular). Stylodia (styluli) (one to) three (or four), free or more or less connate. Stigmas usually erect or reflexed (rarely indistinct), papillate, Dry type. Male flowers often with pistillodia.

Ovules Placentation subbasal, lateral or subapical. Ovules usually one per carpel (rarely one per ovary), anatropous, hemianatropous, campylotropous or orthotropous, apotropous, bitegmic, crassinucellar. Micropyle exostomal, endostomal or bistomal. Outer integument usually eight or more cell layers thick (in Nypa ten cell layers thick). Inner integument usually two or three (rarely up to seven) cell layers thick. Parietal cell formed from archesporial cell. Parietal tissue one to six cell layers thick (sometimes absent). Nucellar cap often formed by periclinal divisions of megasporangial epidermis. Megagametophyte usually monosporous, Polygonum type (rarely disporous, Allium type). Antipodal cells persistent, often proliferating. Endosperm development ab initio nuclear. Endosperm haustorium micropylar. Embryogenesis onagrad or asterad. Polyembryony occurring in some species.

Fruit A usually single-seeded (rarely two- to ten-seeded; sometimes dry) drupe (sometimes with seed and endocarp almost fused; in some species very large: Lodoicea having seeds with a length of up to 50 cm and a weight of up to 18 kg) or a berry (sometimes dry); often with persistent style and/or stigmas.

Seeds Aril sometimes present. Sarcotesta sometimes present. Testa usually with two outer layers thickened. Tegmen? Perisperm not developed. Endosperm copious, often ruminate, with aleurone, lipids and hemicellulose; starch absent; endosperm cells thick-walled. Embryo short, wide, without chlorophyll. Cotyledon one, not photosynthesizing. Cotyledon hyperphyll elongate to compact, not assimilating. Hypocotyl internode and mesocotyl absent. Coleoptile present or absent. Radicula stout, branched, persistent or ephemeral, with short collar (sometimes with roots). Germination cryptocotylar.

Cytology n = 13–18 (rarely more); n = 13, 14 (Calamoideae), n = 17? (Nypa), n = 18 (Coryphoideae), n =13 (Ceroxyloideae), n = 16 (Arecoideae) – Polyploidy rarely occurring (e.g. n = 303±3 in Voaniola).

DNA

Phytochemistry Flavone-C-glycosides, flavone-5-glycosides, flavone sulfate, flavonol sulfate, luteolin sulfate, cyanidin, tricine (tricetine-3’,5’-dimethyl ether; frequent), diterpenes, sesquiterpenes, tannins, catechins, proanthocyanidins, triterpene saponins, or steroidal saponins present. Flavonols (kaempferol, quercetin, luteolin, etc.) and pyrimidine alkaloids rare. Ellagic acid and cyanogenic compounds not found. Lignins with p-coumarylic alcohol and coniferyl and sinapyl monomers.

Use Ornamental plants, fruits (Cocos, Phoenix, Salacca), vegetables (Euterpe etc.), starch sources (Metroxylon), sugar and alcohol (Arenga, Borassus, Caryota, Phoenix), stimulants (Areca catechu), medicinal plants, fruit-oils and fats (Cocos, Elaeis, Orbigyna), waxes (Ceroxylon, Copernicia), fibres for roofing, textiles and carpets (Cocos), buttons and carvings (seeds of Phytelephas), tools, weapons, basketry, handicraft, timber, carpentries, jewellery, perfumes.

Systematics The subdivision below is founded on Dransfield & al. (2005) and the strict consensus tree in Baker & al. (2009). Calamoideae are sister to all other Arecaceae. A weakly supported sister-group relationship between Arecaceae and Dasypogonaceae was identified by Barrett & al. (2013).

Calamoideae Beilschm. in Flora 16(Beibl. 7): 55, 105. 14 Jun 1833 [‘Calameae’]

21/c 650. Pantropical, with their highest diversity in tropical Asia. Root periderm absent. Endodermal cells walls usually thickened. Leaves with reduplicate ptyxis. Adaxial subepidermal non-vascular fibres present. Longitudinal bundles bridging to adaxial epidermis via vertically elongate sclereids. Lateral vascular bundles adaxial to longitudinal bundles. Parenchyma cells adjacent to protoxylem usually inflated. Epidermal cells rectangular, with sinuate anticlinal walls. Flowers inserted in dyads. Tepals usually connate. Inner tepals with valvate aestivation. Pollen grains in Calameae equatorially disulcate. Ovary and fruit covered with reflexed scales. Stylodia usually separate (sometimes absent). Placentation basal. Ovule probably epitropous, with spirally twisted funicle. Fruit enclosed by reflexed scales. Endocarp usually thin. Seeds one to three. Sarcotesta usually thick. n = 13, 14. – Ancistrophyllinae are sister-group to the remaining Calamoideae.

Ancistrophyllinae Becc. in Ann. Roy. Bot. Gard. Calcutta 12(2): 209. 1918 [‘Ancistrophyllae’]

3/22. Oncocalamus (5; tropical Africa), Laccosperma (5; tropical Africa), Eremospatha (12; tropical Africa). – Tropical Africa.

[Lepidocaryeae+Calameae]

Lepidocaryeae (Mart.) Dumort., Anal. Fam. Plant.: 55. 1829

5/c 32. Eugeissoninae Becc. in Ann. Roy. Bot. Gard. Calcutta 12(2–3): 210. 1918 [‘Eugeissoneae’]. Eugeissona (6; Thailand, the Malay Peninsula, Borneo), Raphia (c 20; tropical and southern Africa, Madagascar, one species from Nicaragua to Brazil). – Mauritiinae Meisn., Plant. Vasc. Gen.: Tab. Diagn. 354, Comm. 265. 13-15 Feb 1842 [‘Mauritieae’]. Mauritia (2; tropical South America), Mauritiella (3; northern South America), Lepidocaryum (1; L. tenue; western Amazonia). – Pantropical.

Calameae (Kunth) Lecoq et Juill., Dict. Rais. Term. Bot.: 98. 1831

13/c 600. Korthalsiinae Becc. in Ann. Roy. Bot. Gard. Calcutta 12(2): 209. 1918 [‘Korthalsieae’]. Korthalsia (26; tropical Asia), Eleiodoxa (1; E. conferta; West Malesia), Salacca (c 20; tropical Asia). – Metroxylinae Blume in Rumphia 2: 176. Jan-Aug 1843 [‘Metroxyleae’]. Metroxylon (c 8; East Malesia, the Solomon Islands, Vanuatu, Fiji, Samoa), Pigafetta (2; Central and East Malesia). – Plectocomiinae J. Dransf. et N. W. Uhl in Principes 30: 5. 20 Feb 1986. Myrialepis (1; M. paradoxa; the Malay Peninsula, Sumatra), Plectocomiopsis (5; southern Burma, Thailand, West Malesia), Plectocomia (16; tropical Asia to West Malesia). – Calaminae Meisn., Plant. Vasc. Gen.: Tab. Diagn. 354, Comm. 265. 13-15 Feb 1842 [‘Calameae’]. Calamus (c 400; tropical Asia, Queensland, the Solomon Islands, Vanuatu, Fiji, one species, C. deerratus, in tropical West and Central Africa, with their highest diversity in Malesia), Ceratolobus (6; West Malesia); Daemonorops (c 115; tropical Asia, with their largest diversity in West Malesia), Pogonotium (3; West Malesia), Retispatha (1; R. dumetosa; Borneo). – Tropical Africa, tropical Asia to Samoa, with their largest diversity in West Malesia. Pollen grains equatorially disulcate. – Korthalsiinae are sister-group to the remaining Calameae, and Metroxylinae are sister to the clade [Plectocomiinae+Calaminae]. Calameae are usually climbing by a cirrhus, a prolonged hooked leaf apex and often by a flagellum produced by the spiny modified inflorescence axis, which is adnate to the leaf sheath at the nearest node above.

[Nypa+[Coryphoideae+[Ceroxyloideae+Arecoideae]]]

Stevens (2001 onwards) lists the following features common to this clade: primary growth sustained; presence of root periderm; straight anticlinal epidermal walls; and absence of adaxial subepidermal non-vascular fibres.

Nypoideae Griff., Palms Brit. E. Ind.: 7. post 3 Sep 1850 [‘Nipinae’]

1/1. Nypa (1; N. fruticans; Sri Lanka, Bengal, Southeast Asia, Malesia, Melanesia, northern Queensland). – Fossils of Nypa are known from the Late Cretaceous and the Early Paleogene of many parts of the world (e.g. Tasmania and the London Clay flora) and indicate an almost worldwide former distribution. Branching dichotomous. Endodermal cell walls not or only little thickened. Silica bodies small, hat-shaped. Leaves pseudopinnate, with reduplicate ptyxis. Veins sinuate, irregular, bridging to epidermis via vertically elongated sclereids. Sheaths of transverse veins sclereidal. Hypodermal cells several layered, with lignified walls, hexagonal, transversely elongate. Epidermal cells hexagonal to fusiform. Hydathodes present. Stomatal guard cells in cross-section with several ledges. Inflorescence axis adnate to internode above. Male inflorescence spike. Female inflorescence capitate. Inner tepals free. Tepals in male flowers 3+3, undifferentiated. Stamens three, antesepalous. Filaments connate. Anthers extrorse. Staminodia absent. Pollen grains zonasulcate (sulcus encircling). Tepals in female flowers absent? Pistil composed of three (or four) connate carpels with conduplicate margins. Pistillodium absent. Placentation laminar to submarginal. Outer integument approx. ten cell layers thick. n = 17? – Nypa fruticans, the sole extant species of this clade, is sister to the remaining Arecaceae.

[Coryphoideae+[Ceroxyloideae+Arecoideae]]

The following potential synapomorphies are listed by Stevens (2001 onwards): sieve tubes with compound sieve plates; cell walls of endodermis with U-shaped thickenings; and microsporogenesis simultaneous.

Coryphoideae Burnett, Outlines Bot.: 398. Feb 1835 [‘Coryphidae’]

42/455–465. Tropical, subtropical and warm temperate regions in both hemispheres, with fewer species in South America. Stem usually unbranched. Metaxylem elements in transverse section often more than two. Leaves usually pseudopalmate or costapalmate, with induplicate ptyxis. Mesophyll without free fibre bundles. Longitudinal bundles with adaxially-abaxially elongate bridging sclereids. transverse bundles with broad sheath of fibres. Adaxial vein rib with five or more independent vascular bundles. Inflorescence varying in shape. Inner tepals often with valvate aestivation, usually connate. Microsporocyte usually with ring of callose. Carpels usually free or with connate stylodia (style sometimes present and with three connate or one stylar canal; style sometimes absent). n = 18.

[Chuniophoeniceae+[Corypha+[Caryoteae+Borasseae]]]

This clade is sister-group to the remaining Coryphoideae in Baker & al. (2009).

Chuniophoeniceae J. Dransf., N. W. Uhl, Rasmussen, W. J. Baker, M. M. Harley et C. Lewis in Kew Bull. 60: 561. 2006

3/4. Chuniophoenix (2; southern China, Hainan, Vietnam), Kerriodoxa (1; K. elegans; peninsular Thailand), Nannorrhops (1; N. ritchieana; the Arabian Peninsula to Pakistan). – The Arabian Peninsula and eastwards to Vietnam. – Chuniophoeniceae are sister to a clade comprising [Corypha+[Caryoteae+ Borasseae]].

[Corypha+[Caryoteae+Borasseae]]

Corypheae Martinov, Tekhno-Bot. Slovar: 166. 3 Aug 1820 [‘Coryphineae’]

1/6–8. Corypha (6–8; India, Sri Lanka, Southeast Asia, Malesia, Queensland).

[Caryoteae+Borasseae]

Caryoteae Scheff. in Ann. Jard. Bot. Buitenzorg 1: 142. 1876

3/36. Caryota (12; India, Sri Lanka, Southeast Asia, Malesia, tropical Australia, the Solomon Islands, Vanuatu), Arenga (c 17; Himalayas, southern China, Taiwan, Ryukyu Islands, Malesia, Queensland), Wallichia (7; eastern Himalayas, southern China). – Tropical Asia, tropical Australia, Melanesia. Style absent.

Borasseae Dumort., Anal. Fam. Plant.: 55. 1829. Bismarckia (1), Satranala (1), 6/20–25. Hyphaene (c 10; Africa, Madagascar, the Mascarene Islands, the Arabian Peninsula, India), Medemia (1; M. argun; northern tropical Africa), Latania (3; Mauritius, Réunion), Lodoicea (1; L. maldivica; the Seychelles), Borassodendron (2; the Malay Peninsula, Borneo), Borassus (5–7; tropical and southern Africa, Madagascar, Sri Lanka to Malesia). – Tropical regions in the Old World. – Lodoicea maldivica has the largest seed among all seed-plants, weighing up to 15–20(–30) kg and having a length of up to at least 4 dm.

[Phoenix+[Sabal+Cryosophileae]]

This clade is sister-group to Trachycarpeae.

Phoeniceae Horan., Char. Ess. Fam.: 43. 17 Jun 1847

1/13. Phoenix (13; Macaronesia, the Mediterranean, tropical and subtropical regions in Africa and Asia eastwards to the Malay Peninsula and Sumatra).

[Sabal+Cryosophileae]

Sabaleae (Martius) Dumort., Anal. Fam. Plant.: 55. 1829

1/16. Sabal (16; southeastern United States to South America).

Cryosophileae J. Dransf., N. W. Uhl, Rasmussen, W. J. Baker, M. M. Harley et C. Lewis in Kew Bull. 60: 561. 2006

9/c 77. Schippia (1; S. concolor; Belize), Trithrinax (3; Brazil, Bolivia, Paraguay, Uruguay, Argentina), Zombia (1; Z. antillarum; Hispaniola), Coccothrinax (c 50; the West Indies, with their highest diversity on Cuba), Hemithrinax (2; Cuba), Thrinax (5; the West Indies), Chelyocarpus (4; tropical South America), Cryosophila (10; western Mexico to northern Colombia), Itaya (1; I. amicorum; Brazil, Peru), Sabinaria (1; S. magnifica; southern Panamá, northwestern Colombia). – Tropical America.

Trachycarpeae Satake in Hikobia 3: 121. Oct 1962

18/c 285. Washingtonia (2; arid and semiarid regions in southwestern North America), Pritchardia (35; the Solomon Islands, Fiji, Samoa, Tonga, the Cook Islands, the Tuamotu Islands, the Hawaiian Islands), Copernicia (≤25; the West Indies, South America, with their largest diversity on Cuba), Colpothrinax (3; Central America, Cuba), Brahea (12; Central America), Rhapidophyllum (1; R. hystrix; southeastern United States), Maxburretia (3; Thailand, the Malay Peninsula), Chamaerops (1; C. humilis; western Mediterranean), Trachycarpus (7; Himalayas, China, northern Thailand), Guihaia (2; southern China, northern Vietnam), Rhapis (8; southern China, Southeast Asia, West Malesia), Serenoa (1; S. repens; southeastern United States), Acoelorraphe (1; A. wrightii; Central America), Livistona (33; northeastern Africa, the Arabian Peninsula, Ryukyu Islands, tropical Asia, tropical Australia), Johannesteijsmannia (13; Hainan, Vietnam, West Malesia), Pholidocarpus (6; Malesia), Saribus (4; the Philippines, New Guinea, Nggela in the Solomon Islands, southernmost New Caledonia), Licuala (c 125; India, Southeast Asia, Malesia, tropical Australia, Vanuatu). – Nearly pantropical. – Colpothrinax is sister to the remaining Trachycarpeae.

[Ceroxyloideae+Arecoideae]

According to Stevens (2001 onwards) Ceroxyloideae and Arecoideae have the following synapomorphies in common: petiole vascular bundles arranged in one or more V’s; sheaths of transverse veins sclereidal; veins sinuate, irregular; and epidermal cells hexagonal to fusiform.

Ceroxyloideae Drude in Bot. Zeitung (Berlin) 35: 632. 28 Sep 1877 [‘Ceroxylinae’]

8/43. Madagascar, the Comoro Islands, northeastern Queensland, Florida, Central America, the West Indies, northern South America. Leaves pseudopinnate, with reduplicate ptyxis. Flowers solitary along inflorescence rhachis. Tepals sometimes elongate. n = 13. – Phytelephas and its closest relatives have tetramerous flowers with up to c. 1.000 centrifugal stamens (Palandra) and approx. ten carpels. Ceroxyloideae are sister-group to Arecoideae and Cyclospatheae are sister to the clade [Ceroxyleae+Phytelephanteae] (Baker & al. 2009).

Cyclospatheae O. F. Cook in A. P. Northrop, Mem. Torrey Bot. Club 12: 24. 1902

1/4. Pseudophoenix (4; Florida, Central America, the West Indies).

[Ceroxyleae+Phytelephanteae]

Ceroxyleae Blatter, Palms Brit. India Ceylon: xvi, 337. 1926 [‘Ceroxylinae’]

4/30. Ceroxylon (11; the Andes), Juania (1; J. australis; Juan Fernández Islands); Oraniopsis (1; O. appendiculata; northeastern Queensland), Ravenea (17; Madagascar, the Comoro Islands). – Madagascar, the Comoro Islands, Queensland, western South America.

Phytelephanteae Horan., Char. Ess. Fam.: 38. 17 Jun 1847

3/9. Phytelephas (6; southern Panamá, Colombia, Ecuador, Peru, Bolivia), Aphandra (1; A. natalia; eastern Ecuador), Ammandra (2; western and eastern Colombia). – Northern South America.

Arecoideae Burnett, Outlines Bot.: 401. Feb 1835 [‘Arecidae’]

c 110/1.270–1.340. Pantropical. Stem with crownshaft formed by elongated leaf sheaths. Leaves with reduplicate ptyxis. Hypodermal cells hexagonal, transversely elongate. Inflorescence with prophylls and one or several bracts. Flowers usually in triads, when monoecious with central female flower and lateral male flowers, or in acervuli, with flowers in two vertical rows. Stylodia usually separate (style sometimes single). n = 16. – Iriarteeae are sister-group of the remaining Arecoideae. The apical meristem in the aerial stem in Iriarteeae increases its size, the stem gradually growing thicker (stilt roots arising from the lower part of the stem stabilize the otherwise very unstable structure of the plant). The flowers in Chamaedoreae are often sessile and arranged in acervuli, modified often ebracteate cincinni.

Iriarteeae Drude in Bot. Zeitung (Berlin) 35: 632. 28 Sep 1877 [‘Iriarteae’]

5/32. Iriartea (1; I. deltoidea; Central America, tropical South America), Socratea (5; northern South America), Dictyocaryum (3; tropical South America), Iriartella (2; tropical South America), Wettinia (21; tropical South America). – Central America, tropical South America.

[Chamaedoreeae+[[Roystonea+[Reinhardtia+Cocoseae]]+[[Podococcus+[Orania+Sclerosperma]]+[Euterpeae+[Pelagodoxeae+[Leopoldinia+[Manicaria+Geonomateae]]]+Areceae]]]]

Chamaedoreeae Drude in Bot. Zeitung (Berlin) 35: 632. 28 Sep 1877 [‘Chamaedorineae’]

5/90–125. Hyophorbe (5; Mauritius, Réunion), Synechanthus (2; Mexico, Central America, tropical South America), Gaussia (5; Central America, the West Indies), Chamaedorea (77–>110; tropical America), Wendlandiella (1; W. gracilis; Brazil, Peru). – The Mascarene Islands, tropical America. – Chamaedoreeae are sister-group to the remaining Arecoideae.

[[Roystonea+[Reinhardtia+Cocoseae]]+[[Podococcus+[Orania+Sclerosperma]]+[Euterpeae+[Pelagodoxeae+[Leopoldinia+[Manicaria+Geonomateae]]]+Areceae]]]

Roystoneeae J. Dransf., N. W. Uhl, Rasmussen, W. J. Baker, M. M. Harley et C. Lewis in Kew Bull. 60: 562. 2006 [‘Roystoneae’]

1/10. Roystonea (10; the West Indies, northeastern South America).

Reinhardtieae J. Dransf., N. W. Uhl, Rasmussen, W. J. Baker, M. M. Harley et C. Lewis in Kew Bull. 60: 562. 2006

1/6. Reinhardtia (6; Central America).

Cocoseae Dumort., Anal. Fam. Plant.: 56. 1829 [‘Cocoineae’]

19/290–305. Elaeidinae Drude in C. F. P. von Martius, Fl. Bras. 3(2): 395. 1 Mai 1882 [‘Elaeideae’]. Barcella (1; B. odora; Brazil), Elaeis (2; E. guineensis: tropical West and Southwest Africa from Gambia to Angola; E. oleifera: Central America, tropical South America). – Bactridinae Drude in C. F. P. von Martius, Fl. Bras. 3(2): 301, 395. 1 Mai 1882. Acrocomia (4; Central America, Cuba, tropical South America), Aiphanes (22; tropical America), Astrocaryum (c 35; tropical America), Bactris (c 75; tropical America), Desmoncus (24; tropical America). – Attaleinae Drude in C. F. P. von Martius, Fl. Bras. 3(2): 395. 1 Mai 1882 [‘Attaleeae’]. Beccariophoenix (1; B. madagascariensis; eastern Madagascar), Jubaeopsis (1; J. caffra; Pondoland in Eastern Cape), ‘Butia’ (11; South America; non-monophyletic), Jubaea (1; J. chilensis; 32°S to 35°S in central Chile), Paschalococos (1; P. disperta; Easter Island, extinct), Cocos (1; C. nucifera; tropical and subtropical coastal areas), ‘Syagrus’ (32; tropical South America, one species in the eastern West Indies; polyphyletic), Parajubaea (3; subalpine regions in Colombia, Ecuador and Bolivia), Attalea (72; tropical America), Lytocaryum (2; southeastern Brazil), Voanioala (1; V. gerardii; Madagascar), Allagoptera (5; Brazil, Bolivia, Paraguay, Argentina). – Tropical and subtropical Africa and America, Madagascar. – Roystonea is sister to the clade [Reinhardtia+Cocoseae]. Elaeidinae are sister to Bactridinae, and Beccariophoenix is sister to the remaining Attaleinae. Roystoneeae are sister-group to the rest of Arecoideae.

[[Podococcus+[Orania+Sclerosperma]]+[Euterpeae+[Pelagodoxeae+[Leopoldinia+[Manicaria+Geonomateae]]]+Areceae]]

Podococceae J. Dransf. et N. W. Uhl in Principes 30: 6. 20 Feb 1986

1/1. Podococcus (1; P. barteri; tropical Africa). – Podococcus is sister to [Orania+Sclerosperma], and Podococceae are sister-group to a trichotomy comprising the remaining Arecoideae.

[Orania+Sclerosperma]

Oranieae Becc. in O. Beccari et R. E. G. Pichi Sermolli, Webbia 11: 15. 1955

1/28. Orania (28; Madagascar, southern Thailand and eastwards to New Guinea, tropical Australia).

Sclerospermateae J. Dransf., N. W. Uhl, Rasmussen, W. J. Baker, M. M. Harley et C. Lewis in Kew Bull. 60: 562. 2006 [‘Sclerospermeae’]

1/3. Sclerosperma (3; tropical West Africa).

[Euterpeae+[Pelagodoxeae+[Leopoldinia+[Manicaria+Geonomateae]]]+Areceae]

Euterpeae J. Dransf., N. W. Uhl, Rasmussen, W. J. Baker, M. M. Harley et C. Lewis in Kew Bull. 60: 562. 2006

5/53. Hyospathe (2; tropical America), Euterpe (30; tropical America), Prestoea (11; Central America), Neonicholsonia (1; N. watsonii; Central America), Oenocarpus (9; tropical South America). – Tropical America. – Hyospathe is sister to the rest of Euterpeae, although with fairly low bootstrap support.

Pelagodoxeae+[Leopoldinia+[Manicaria+Geonomateae]]]+Areceae]

Pelagodoxeae J. Dransf., N. W. Uhl, Rasmussen, W. J. Baker, M. M. Harley et C. Lewis in Kew Bull. 60: 563. 2006

2/2–4. Pelagodoxa (1; P. henryana; Melanesia, the Marquesas Islands), Sommieria (1–3; S. leucophylla; New Guinea). – New Guinea, Melanesia, the Marquesas Islands. – Pelagodoxeae are sister to the clade [Leopoldinia+[Manicaria+Geonomateae]].

[Leopoldinia+[Manicaria+Geonomateae]]

Leopoldinieae J. Dransf., N. W. Uhl, Rasmussen, W. J. Baker, M. M. Harley et C. Lewis in Kew Bull. 60: 562. 2006

1/3. Leopoldinia (3; Colombia, Venezuela, Amazonian Brazil).

[Manicaria+Geonomateae]

Manicarieae J. Dransf., N. W. Uhl, Rasmussen, W. J. Baker, M. M. Harley et C. Lewis in Kew Bull. 60: 562. 2006

1/1. Manicaria (1; M. saccifera; Central America, Trinidad, tropical South America).

Geonomateae Luerss., Handb. Syst. Bot. 2: 342. Jan 1880 [‘Geonomeae’]

6/c 105. Welfia (2; Nicaragua to Colombia and Ecuador, central Peru), Pholidostachys (7; Nicaragua to Colombia, Ecuador, Peru, Brazil); Calyptrogyne (18; Central America), Calyptronoma (3; the West Indies), Asterogyne (5; tropical America), Geonoma (68; tropical America). – Tropical America. – The [Welfia+Pholidostachys] clade is sister to the remaining Geonomateae.

Areceae (Mart.) Dumort., Anal. Fam. Plant.: 56. 1829 [‘Arecaceae’]. c 60/650–690. – Dypsidinae are sister-group to the remaining Areceae.

Dypsidinae Becc., Palmae Madagascar: 2. 1912 [‘Dypsideae’]

3/140–145. Lemurophoenix (1; L. halleuxii; northeastern Madagascar), Dypsis (c 140; Madagascar, the Comoro Islands, Pemba), Marojejya (2; northeastern Madagascar). – Madagascar, the Comoro Islands, Pemba.

[Arecinae+[ODAT clade+Iguanurinae]+Ptychospermatinae]

Arecinae Engl. in Bopt. Jahresber. (Just) 3: 456. Mai-Dec 1877

6/220–225. Cyrtostachys (7; Malesia to Melanesia), Bentinckia (2; India, the Nicobar Islands), Clinostigma (11; New Britain, Vanuatu, Fiji, Samoa, Micronesia to Bonin Island), Areca (65–70; tropical Asia), Nenga (5; Southeast Asia, Malesia), Pinanga (135–140; India and Nepal to southern China, Southeast Asia, Malesia to New Guinea). – Tropical Asia, Melanesia, Samoa, Micronesia. – Arecinae are sister to a clade comprising the “ODAT clade” plus an extended Iguanurinae.

The “ODAT clade”

4/8–10. Oncosperma (5; Sri Lanka, Indochina, Malesia to the Moluccas and the Philippines), Deckenia (1; D. nobilis; the Seychelles), Acanthophoenix (1–3; A. rubra; the Mascarene Islands), Tectiphiala (1; T. ferox; Mauritius). – Tropical Asia, the Mascarene Islands, the Seychelles.

Iguanurinae Benth. et Hook. f., Gen. Plant. 3: 872, 876. 14 Apr 1883 [‘Iguanureae’]

8/c 38. Dictyosperma (1; D. album; the Mascarene Islands), Rhopaloblaste (6; the Nicobar Islands, Malesia and eastwards to the Solomon Islands); Masoala (2; northeastern Madagascar), Iguanura (c 25; West Malesia), Verschaffeltia (1; V. splendida; the Seychelles), Roscheria (1; R. melanochaetes; the Seychelles), Phoenicophorium (1; P. borsigianum; the Seychelles), Nephrosperma (1; N. vanhoutteanum; the Seychelles). – Madagascar, the Mascarene Islands, the Seychelles, the Nicobar Islands, Malesia, the Solomon Islands. – The clade [Dictyosperma+ Rhopaloblaste] is sister to the remaining Iguanurinae.

Ptychospermatinae Benth. et Hook. f., Gen. Plant. 3: 872, 874. 14 Apr 1883 [‘Ptychospermeae’]

38/244–270. Lepidorrhachis (1; L. mooreana; Lord Howe Island); Howea (2; Lord Howe Island); Calyptrocalyx (c 25; the Moluccas, New Guinea); Hydriastele (48–52; East Malesia to Queensland and Fiji), Loxococcus (1; L. rupicola; Sri Lanka); Hedyscepe (1; H. canterburyana; Lord Howe Island), Rhopalostylis (2–3; New Zealand, Chatham Islands, Norfolk Island, Kermadec Islands); Carpoxylon (1; C. macrospermum; Vanuatu), Satakentia (1; S. liukiuensis; Ryukyu Islands), Neoveitchia (1; N. storckii; Vanuatu, Fiji); Actinorhytis (2; New Guinea); Archontophoenix (6; eastern Queensland), Chambeyronia (2; New Caledonia), Actinokentia (2; New Caledonia), Kentiopsis (4; New Caledonia); Cyphokentia (1; C. macrostachya; New Caledonia), Clinosperma (4; New Caledonia); Dransfieldia (1; D. micrantha; western New Guinea), Linospadix (7–11; New Guinea, eastern Queensland, northeastern New South Wales), Laccospadix (1; L. australasica; northeastern Queensland), Heterospathe (c 40; Central and East Malesia, New Guinea, Fiji); Physokentia (7; New Britain, the Solomon Islands, Vanuatu, Fiji), Cyphosperma (4; New Caledonia, Fiji), Burretiokentia (5; New Caledonia), Basselinia (11; New Caledonia), Cyphophoenix (2; New Caledonia). – Ptychospermatinae form a polytomy in the strict consensus tree in Baker & al. (2009; see also Baker & al. 2011 and Alapetite & al. 2014), and have received a much wider definition here than in Dransfield & al. (2005). The following clade received high bootstrap support: Ptychosperma (27; East Malesia, New Guinea, tropical Australia), Normanbya (1; N. normanbyi; New Guinea, northeastern Queensland), Carpentaria (1; C. acuminata; tropical Australia), Wodyetia (1; W. bifurcata; Melville Range in Queensland), Ponapea (3; the Caroline Islands), Drymophloeus (7–15; the Moluccas and eastwards to Samoa), Brassiophoenix (2; New Guinea), Ptychococcus (2–7; New Guinea, the Solomon Islands), Ponapea (3; the Caroline Islands), Wallaceodoxa (1; W. raja-ampat; Gag Island and Waigeo Island off the western peninsula of New Guinea), Adonidia (1; A. merrillii; the Philippines), Jailoloa (1; J. halmaherensis; Halmahera in the Moluccas), Manjekia (1; M. maturbongsii; Biak Island Northwest of New Guinea), Solfia (1; S. samoensis; Samoa), Balaka (7–11; Fiji, Samoa), Veitchia (8; Palawan, Vanuatu, Fiji, Tonga). – Sri Lanka, Ryukyu Islands, North, Central and East Malesia, northeastern Australia, Lord Howe Island, Melanesia, New Zealand and surrounding islands, Tonga, Samoa.

Phylogeny (simplified) of Arecaceae based on morphological and molecular data (Baker & al. 2009).


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