PROTEALES Dumortier Main Tree, Synapomorphies

Proteanae Takht., Sist. Filog. Cvetk. Rast. [Syst. Phylog. Magnolioph.]: 401. 4 Feb 1967

Habit Usually bisexual (sometimes monoecious, andromonoecious or dioecious), evergreen trees or usually shrubs (rarely perennial rhizomatous herbs, some species with lignotuber). The majority are xerophytes. A few species are aquatic.

Vegetative anatomy Mycorrhiza often absent. Phellogen usually ab initio superficial (rarely deep or absent). Secondary lateral growth usually normal (rarely absent). Vessel elements usually with simple (sometimes scalariform or reticulate) perforation plates; lateral pits scalariform, alternate or opposite, simple or bordered pits. Vestured pits sometimes present. Imperforate tracheary xylem elements tracheids, fibre tracheids or libriform fibres with simple and/or bordered pits, septate or non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, usually homocellular (sometimes heterocellular; rarely absent). Axial parenchyma usually paratracheal scanty, vasicentric, aliform, lozenge-aliform, winged-aliform, confluent, scalariform, unilateral, or banded (sometimes apotracheal diffuse or diffuse-in-aggregates; rarely absent). Sieve tube plastids S type (rarely Ss type). Nodes usually 3:3, trilacunar with three leaf traces (sometimes 1:1, 5:5 or 7:7). Secretory cavities absent. Tanniniferous cells often abundant. Heartwood with gum-like substances. Sclereids usually numerous. Silica bodies present in some species. Prismatic calciumoxalate crystals sometimes present.

Trichomes Hairs unicellular or multicellular (usually tricellular), uniseriate and thick-walled, basal cell being sunken into epidermis, stalk cell short and terminal cell long and sometimes split (rarely candelabra-shaped, dendritic or almost stellate), or absent; glandular hairs rare.

Leaves Usually alternate (spiral or rarely distichous; sometimes opposite or verticillate), simple or pinnately compound (sometimes twice or several times), usually entire or pinnately lobed (rarely palmate or palmately compound), usually xeromorphic (thick, coriaceous, ericoid or hair-like), with usually conduplicate (sometimes plicate) ptyxis. Stipules usually absent (rarely pairwise or single, sheathing, with ocreate base); leaf sheath usually absent. Petiole vascular bundle transection at least sometimes annular. Venation usually pinnate, brochidodromous (rarely parallellodromous or palmate), or leaves one-veined. Stomata usually (brachy)paracytic (rarely anomocytic or laterocytic). Cuticular wax crystalloids as clustered tubuli (Berberis type), chemically usually dominated by secondary alcohol nonacosan-10-ol (in Nelumbo -4-10- or -5-10-diols). Secretory cavities usually absent (rarely with laticifers). Mesophyll usually with sclerenchymatous idioblasts (with asterosclereids, columellar or palisade sclereids). Idioblasts with ethereal oils absent. Leaf margin serrate (sometimes serrate-dentate), crenate or entire.

Inflorescence Flowers solitary or pairwise, usually in terminal or axillary spike-, spadix-, cone-, or head-like, or compound inflorescence (flowers rarely solitary axillary or in compact multiflorous globular inflorescence). Numerous species with pseudanthia surrounded by showy involucral bracts.

Flowers Usually zygomorphic (rarely actinomorphic or asymmetric). Hypanthium present or absent. Hypogyny or half epigyny. Sepals (three or) four (to seven), free, with valvate aestivation, usually petaloid, free or connate into tube (tepals rarely numerous, with imbricate aestivation, outer ones sepaloid, inner ones petaloid, spiral). Small, almost spiny, abaxial projection present immediately below sepal apex. Petals probably absent; in their place (two to) four hypogynous alternisepalous scales or glands (= rudimentary petals?), free or connate into annular extrastaminal nectariferous disc (nectary and disc absent in some species).

Androecium Stamens (three or) four (to more than 400), usually whorled (rarely spiral), haplostemonous, antesepalous. Filaments usually free, entirely or in part adnate to sepals (episepalous, sometimes free from tepals; rarely absent). Anthers usually free, basifixed, non-versatile, usually tetrasporangiate (sometimes disporangiate), usually introrse (rarely latrorse or extrorse), longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia usually absent (female flowers sometimes with three or four staminodia).

Pollen grains Microsporogenesis simultaneous. Pollen grains usually triporate (sometimes di- to octoporate, or di- to octocolporate; rarely di- to octocolpate, tricolpoidate, rugate, monocolpate or spiraperturate), shed as monads, usually bicellular (rarely tricellular) at dispersal. Apertures arranged according to Garside’s rule. Exine tectate or semitectate, with columellate infratectum, reticulate, perforate or rugulate, scabrate or echinate.

Gynoecium Pistil usually composed of a single, entirely or partially closed carpel, usually with gynophore; carpel usually plicate (rarely ascidiate with margins seemingly occluded by secretion), postgenitally entirely fused, without canal (carpels rarely two to c. 40, whorled, free). Ovary superior or semi-inferior, usually monocarpellate and unilocular. Stylodium usually single (rarely absent or three to nine), usually with apex expanded around stigma and specialized for secondary pollen presentation. Stigma usually terminal, subterminal or lateral (stigmas rarely decurrent on adaxial side of each stylodium), usually papillate, Dry (rarely Wet) type. Pistillodium usually absent (male flowers rarely with pistillodium).

Ovules Placentation marginal eller subapical (rarely apical or ventral). Ovules one to three (to more than 100) per carpel, usually hemitropous (sometimes anatropous, amphitropous or orthotropous), pendulous, bitegmic, crassinucellar. Micropyle usually endostomal. Inner integument usually thicker than outer integument (not in Nelumbo). Nucellar cap sometimes present. Megagametophyte Polygonum type. Synergids usually with a filiform apparatus. Endosperm formation ab initio nuclear, later usually entirely or partially cellular. Endosperm haustorium chalazal. Embryogenesis usually asterad (rarely solanad).

Fruit A follicle, capsule, achene or drupe, often fused into a cone-shaped syncarp (often lignified and long persistent). Pericarp and integument sometimes fused into a caryopsoid fruit.

Seeds Aril absent. Elaiosome present in some species. Seed coat usually testal. Testa sometimes multiplicative, occasionally thin. Exotesta sometimes palisade. Endotesta often palisade. Exotegmen often fibrous. Endotegmen unspecialized. Perisperm not developed. Endosperm usually absent (sometimes rudimentary, oily and proteinaceous). Embryo large, elongate, straight, oily and proteinaceous (starch absent), well differentiated, with or without chlorophyll. Cotyledons two (to nine). Germination phanerocotylar or cryptocotylar.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), flavonol derivatives, cyanidin, condensed tannins, proanthocyanidins (prodelphinidins), tropane alkaloids (bellendine, methylbellendine, benzoyltropine, benzoyltropane, 2α-benzoyltropane and phenylhydroxymethyl tropane), saponins, sapogenins, naphthoquinones, benzoquinones (rapanone, arbutin), tyrosine-derived cyanogenic glycosides (dhurrin, proteacin, taxiphyllin, triglochinin), and quebrachitol present. Ellagitannins, benzylisoquinoline alkaloids and aporphine alkaloids rare. Ellagic acid not found.

Nelumbo is sister to [Platanus+Proteaceae]. Potential synapomorphies of this clade are, according to Stevens (2001 onwards): woody habit; wood rays at least 8-seriate; stomata laterocytic; flowers tetramerous; stamens equalling tepals, antetepalous; carpels with five vascular bundles, hairy, with complete postgenital fusion; stylodium long; ovules orthotropous; inner integument three to five cell layers thick; endosperm development nuclear; presence of myricetin and non-hydrolyzable tannins; and absence of benzylisoquinoline alkaloids.

Richard in J. B. G. M. Bory de Saint-Vincent, Dict. Class. Hist. Nat. 11: 492. 10 Feb 1827 [’Nelumbiaceae’], nom. cons.

Nelumbonales Bercht. et J. Presl, Přir. Rostlin 1(1): 1. 1823 [‘Nelumbiaceae’]; Nelumbonopsida Endl., Gen. Plant.: 898. Nov 1839 [’Nelumbia’]; Nelumbonanae Takht. ex Reveal in Novon 2: 236. 13 Oct 1992; Nelumbonidae Takht., Divers. Classif. Fl. Pl.: 83. 24 Apr 1997; Nelumbonineae Shipunov in A. Shipunov et J. L. Reveal in Phytotaxa 16: 64. 4 Feb 2011

Distribution East, South and Southeast Asia, Malesia southeastwards to northern Australia, eastern North America to the West Indies and Colombia.

Fossils Fossil vegetative and reproductive structures are known from the Albian of Virginia and from the Campanian to the Maastrichtian of Argentina, and also from Cenozoic localities in the Northern Hemisphere. Peltate leaves have been described under Nelumbo and the similar extinct Nelumbites. Exnelumbites, in the form of leaf macrofossils with glandular chloranthoid teeth and lacking secondary venation, was described from the Campanian-Maastrichtian of Mexico and New Mexico.

Habit Bisexual, perennial herbs. Aquatic. Rhizome rich in starch. Internodes near growth points forming fleshy banana-shaped nutrient-storing tubers.

Vegetative anatomy Mycorrhiza absent. Main root ephemeral and replaced by adventitious roots from nodes. Rhizome with scale-like cataphylls and normal photosynthesizing leaves. Foliar primordia developing in groups of three: one sheathing fleshy scale-like cataphyll enclosing stem (perhaps representing a foliar stipule; at first enclosing apical bud) on lower side of rhizome; one scale-like cataphyll (enclosing petiole base of normal leaf); and one normal leaf on upper side of rhizome. Floral primordium developing in axil of second scale-like cataphyll. Rhizome branch developing in axil of normal leaf. Prophyll adaxial, on axillary branches on same side as first scale-like cataphyll. Phellogen absent? Primary vascular tissue as scattered bundles (atactostele?), without fibrous envelope. Endodermis present. Vessels (protoxylem lacunae?) present in root and rhizome metaxylem. Secondary lateral growth and cambium absent. Primary vessel elements with scalariform perforation plates; lateral pits scalariform, simple pits? Imperforate tracheary xylem elements tracheids with simple? pits. Wood rays absent. Axial parenchyma? Sieve tube plastids Ss type, with approx. ten starch grains unequal in size. Non-dispersive tubular and rod-shaped protein bodies present in sieve elements. Nodes? Parenchyma and vascular tissue with articulated laticifers as thin-walled elongate cells. Sclerenchymatous sclereids absent. Calciumoxalate as single (prismatic?) crystals.

Leaves Alternate (in groups of three along stem, distichous), simple, entire, peltate or sunken, lanceolate, with involute ptyxis. Stipule single, intrapetiolar, sheathing, with ocreate base, open on leaf-opposite side of stem. Petiole up to c. 2 m long, echinate. Petiole vascular bundle transection? Lamina peltate, concave, up to one metre in diameter, usually with central disc responsible for air exchange for petiole canals. Venation palmate, actinodromous; main veins dichotomous, reaching leaf margin. Stomata anomocytic, restricted to adaxial side of leaf (on central disc). Cuticular wax crystalloids as clustered tubuli (Berberis type), chemically dominated by nonacosan-4-10-diol or nonacosan-5-10-diol. Lamina with secretory cavities (laticifers) with latex. Mesophyll without sclerenchymatous idioblasts. Idioblasts with ethereal oils absent. Leaf margin entire.

Inflorescence Flowers axillary, solitary. Floral prophylls (bracteoles) absent.

Flowers Actinomorphic, large. Receptacle massive, with epidermal sharp-pointed calciumoxalate druses. Pedicel up to c. 2 m long. Hypogyny. Tepals numerous; outermost two to five (to eight) tepals sepaloid, with imbricate aestivation, inserted in vertical plane, caducous; inner c. 10 to c. 30 tepals petaloid, with imbricate aestivation, spiral, caducous, free. Nectary absent? Disc absent.

Androecium Stamens c. 100 to c. 300 (to more than 400), spiral (developing from annular meristem). Filaments elongate, free from each other and from tepals. Anthers basifixed, non-versatile, tetrasporangiate, outer stamens extrorse, inner stamens extrorse, introrse or latrorse, longicidal (dehiscing by longitudinal slits); connective with claw-shaped apical appendage. Tapetum secretory, with multinucleate cells. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolpate (rarely monocolpate, dicolpate or spiraperturate), shed as monads, tricellular at dispersal. Exine tectate, with columellate infratectum, rugulate to (micro)perforate, with supratectal processes of various shape.

Gynoecium Carpels (two to) ten to c. 30 (to c. 40), in two to four whorls, free, obconical, fleshy or spongy, sunken into cavities on upper side of receptacle; carpel ascidiate (secondarily), (seemingly) occluded by secretion. Closure by transverse slit occurring together with longitudinal slit. Ovary superior, unilocular (apocarpy). Stylodia absent. Pollen canal central, with elongated papillae. Stigma annularly expanded, papillate, Wet type. Pistillodium absent.

Ovules Placentation ventral to apical (or marginal to apical or laminar-dorsal). Ovule usually one (rarely two) per carpel, anatropous, pendulous, apotropous, bitegmic, crassinucellar. Micropyle endostomal. Outer integument c. 30 cell layers thick. Inner integument eight to ten cell layers thick. Funicular obturator present. Hypostase present. Parietal tissue three to five cell layers thick. Nucellar cap approx. four cell layers thick. Chalaza massive (ovule pachychalazal). Megagametophyte monosporous, Polygonum type. Antipodal cells persistent, proliferating, multinucleate. Endosperm development ab initio cellular? (or helobial? or nuclear?). Endosperm haustorium? chalazal. Embryogenesis solanad (asterad?).

Fruit Hard-walled nuts sunken into dry hard receptacle. Nut with apical pore close to stigmatic remnants. Endocarp fused with seed coat. Pericarp with high concentrations of galactose, mannose and tannins.

Seeds Aril absent. Seeds pachychalazal? Testa thin, indistinct. Mesotesta and endotesta unspecialized. Tegmen unspecialized. Perisperm not developed. Endosperm rudimentary or absent. Embryo large, with chlorophyll. Cotyledons two, fleshy, connate and sheathing into a stipule-like tube surrounding plumule. Radicula ephemeral. Seeds very long-lived (sometimes germinative after at least several hundred years). Germination phanerocotylar.

Phytochemistry Flavonols, flavones, tannins, proanthocyanidins, aporphine and benzylisoquinoline alkaloids (benzoyltetrahydroisoquinoline aporphine, proaporphine, bis-benzoyltetrahydroisoquinoline bases, roemerine, annonaine, liriodenine etc.) present. Ellagic acid, cyanogenic compounds and saponins not found.

Use Ornamental plants, fruits (nut with edible embryo), vegetables. The lotus flower is an important symbol in several oriental religions (‘the Sacred Lotus’).

Platanales T. Lestib. in C. F. P. von Martius, Consp. Regn. Veg.: 12. Sep-Oct 1835 [’Plataneae’]; Platanineae J. Presl in Nowočeská Bibl.. [Wšobecný Rostl.] 7: 1359 [‘1258’], 1379. 1846

Distribution Southeastern Canada to southern Mexico, the Balkan Peninsula, northeastern Mediterranean to western Himalayas, northern Indochina.

Fossils Fossil leaves (simple or palmately to pinnately compound) and reproductive structures of Platanaceae are known from numerous sites in the Northern Hemisphere from the Albian onwards. Fossil leaf taxa include Araliopsoides, Credneria pro parte, Erlingdorfia, Platanites and Tasymia. Aquia brookensis and Friisicarpus brookensis comprise male flowers and infructescences, respectively, from the Albian of Virginia. Hamatia elkneckensis and Friisicarpus elkneckensis are male inflorescences with tricolporate pollen and female flowers and fruits, respectively, from the Early Cenomanian of Maryland. ‘Platananthus’, from the mid-Cretaceous to the Cenozoic of Europe and North America, is represented by male inflorescences bearing flowers with distinct tepals. Quadriplatanus georgianus comprises male and female inflorescences from the Coniacian of Georgia in the United States; the male flowers have four tepals connate at the base; the female flowers bear one outer and one inner whorl of four tepals, the outer whorl being connate into a tube; the eight uniovular carpels are free. Archaranthus krassilovii is a male inflorescence with prominent tepals from the Maastrichtian to the Paleocene of eastern Siberia. Many early Platanaceae were probably insect-pollinated. The diversity and distribution of Platanaceae seem to have reached their peak during the Late Cretaceous and the Palaeogene.

Vegetative anatomy Endomycorrhiza present in at least some species. Phellogen ab initio superficial (later in outer cortex). Vessel elements with scalariform and simple perforation plates; lateral pits alternate, scalariform and/or opposite, simple or bordered pits. Vestured pits present. Imperforate tracheary xylem elements fibre tracheids with simple and bordered pits, septate or non-septate (also vasicentric tracheids). Wood rays multiseriate, homocellular or heterocellular. Axial parenchyma absent or very rare (apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty vasicentric, aliform, confluent, unilateral, or banded). Wood partially storied. Tile cells present. Sieve tube plastids S type. Nodes 7:7, multilacunar with seven leaf traces. Parenchyma with oil cells. Prismatic calciumoxalate crystals often abundant.

Trichomes Hairs unicellular or multicellular, simple or branched, candelabra-shaped, sometimes dendritic or almost stellate; glandular hairs with unicellular globular head present.

Leaves Alternate (usually distichous, sometimes spiral), simple, usually palmately lobed (in Platanus kerrii entire), with plicate ptyxis. Stipules usually pairwise connate, often foliaceous (in Platanus kerrii membranous) and adaxially sheathing (often into tube), usually closed (sometimes open), with ocreate base, early caducous. Axillary bud usually enclosed by petiole base (not in Platanus kerrii). Petiole vascular bundle transection annular, with wing bundles. Venation usually palmate (in Platanus kerrii pinnate); two large secondary veins arising from near leaf base; veins of higher order (tertiary etc.) approaching but not penetrating teeth. Stomata laterocytic or anomocytic. Cuticular wax crystalloids as clustered tubuli (Berberis type), chemically dominated by nonacosan-10-ol. Domatia as pockets in vein axils on lower side of lamina. Idioblasts with ethereal oils absent. Leaf margin usually serrate, with platanoid teeth (glandular with apical cavity; rarely entire).

Inflorescence Axillary, with flowers in one to twelve dense, globular, many-flowered, short-stalked or unstalked partial inflorescences (compact panicles?) on a long pendant common peduncle, each floral head with a single basal circular bract.

Flowers Actinomorphic, small. Hypogyny. Outer tepals three or four (to seven), with valvate aestivation, free or connate at base; odd outer tepal abaxial. Inner tepals usually absent from female flowers (sometimes three or four, rudimentary), in male flowers three or four (to seven), rudimentary, free. Nectary absent. Disc absent.

Androecium Stamens three or four (to seven), antesepalous. Filaments short or absent, free from each other and from tepals. Anthers basifixed, non-versatile, tetrasporangiate, latrorse, longicidal (dehiscing by longitudinal slits); connective expanded into almost peltate terminal appendage. Tapetum probably secretory. Ring of fleshy structures present, possibly representing outer whorl of stamens (‘ridged staminodia’). Female flowers often with three or four staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains tri- or tetracolpate or (hexa)rugate, shed as monads, bicellular at dispersal. Exine semitectate, with columellate infratectum, reticulate.

Gynoecium Carpels (three to) five to eight (or nine), in two (or three) whorls, free; carpel plicate, postgenitally entirely fused, without canal. Ovary superior, unilocular (apocarpy). Stylodia narrowly elongated. Stigmas decurrent in two crests on adaxial side of stylodium, papillate, Dry type. Male flowers often with pistillodium.

Ovules Placentation apical to marginal. Ovule usually one (sometimes two) per carpel, (semi)orthotropous, pendulous, apotropous, bitegmic, crassinucellar. Micropyle endostomal? Outer integument three or four cell layers thick. Inner integument approx. five cell layers thick. Megagametophyte monosporous, Polygonum type? Antipodal cells persistent. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit A globular assemblage (syncarp) of one-seeded achenes, with long hairs at base (fruit rarely follicular).

Seeds Aril absent. Seed coat testal. Testa thin, with hypodermal layer of thickened cells. Mesotesta cells sclerotic. Endotesta unspecialized. Tegmen unspecialized, early disappearing. Perisperm not developed. Endosperm sparse, proteinaceous, oily and with hemicellulose. Embryo large, narrow, straight, well differentiated, chlorophyll? Cotyledons two. Germination phanerocotylar.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), proanthocyanidins (cyanidin, prodelphinidin), and tyrosine derived cyanogenic glycosides (dhurrin, triglochinin) present. Alkaloids, saponins and ellagic acid not found.

Banksiaceae Bercht. et J. Presl, Přir. Rostlin: 235. Jan-Apr 1820 [’Banksiae’]; Proteopsida Bartl., Ord. Nat. Plant.: 91, 110. Sep 1830 [’Proteinae’]; Lepidocarpaceae Schultz Sch., Nat. Syst. Pflanzenr.: 374. 30 Jan-10 Feb 1832; Proteineae Reveal in Kew Bull. 66: 47. Mar 2011

Distribution Africa south of Sahara, Madagascar, southern India and Sri Lanka to eastern China, Taiwan, southern Japan, Indochina, Malesia, islands in the southwestern Pacific, Australia, Tasmania, New Zealand, South and Central America; with their largest diversity in Mediterranean climates in Australia and South Africa.

Fossils Fossils of Proteaceae include wood, leaves, reproductive structures and especially pollen grains from the Santonian of Australia onwards. They have been found in all larger Gondwana fragments, and Late Cretaceous fossils are known also from the Antarctic Peninsula and New Zealand. Macrofossils occur from the Paleocene and the Eocene and include fossilized leaves of Banksieaeformis and Banksieaephyllum and the inflorescence of Musgraveinanthus alcoensis. Reports of Proteaceae fossils from the Northern Hemisphere are questionable.

Vegetative anatomy Mycorrhiza usually absent. Roots with four to seven protoxylem poles, usually with dense bundles of special short superficial lateral roots – proteoid roots, cluster roots – with limited growth (apical meristem aborting), produced at low phosphate supply. Phellogen usually ab initio superficial (rarely deeply seated). Primary medullary rays alternately wide and narrow. Vessel elements usually with simple (rarely scalariform or reticulate) perforation plates; lateral pits scalariform, alternate or opposite, bordered pits. Vestured pits present at least in Persoonia. Imperforate tracheary xylem elements tracheids, fibre tracheids or libriform fibres (sometimes very long) with simple and/or bordered pits, septate or non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, homocellular. Axial parenchyma usually paratracheal scanty, aliform, lozenge-aliform, winged-aliform, confluent, scalariform, vasicentric, unilateral, or banded (rarely apotracheal banded with most bands independent of vessels). Secondary phloem often stratified into hard fibrous and/or sclerenchymatous layers, and soft parenchymatous layers; sieve tubes with sieve surfaces along most of their length. Sieve tube plastids S type (or P type?). Nodes usually 3:3, trilacunar with three leaf traces (sometimes 1:1, unilacunar with one trace; in Finschia trilacunar with three or more traces). Secretory cavities absent. Tanniniferous cells often abundant. Heartwood with resin-like substances. Sclereids abundant. Silica bodies present in some species. Crystals?

Trichomes Hairs tricellular, uniseriate and thick-walled, with basal cell sunken into epidermis, stalk cell short and terminal cell elongate and sometimes bifid, or absent; glandular hairs rare.

Leaves Usually alternate (spiral; sometimes opposite or verticillate), simple or pinnately compound (sometimes twice or several times), usually entire or pinnately lobed (rarely palmate or palmately compound), usually xeromorphic (thick, coriaceous, ericoid or hairy), with usually conduplicate ptyxis. Stipules absent; leaf sheath usually absent. Petiole base often swollen. Petiole vascular bundle transection? Venation usually pinnate, brochidodromous (rarely parallelodromous or palmate), or leaves one-veined. Stomata usually (brachy)paracytic (rarely anomocytic?; in Bellendena laterocytic). Cuticular wax crystalloids as clustered tubuli (Berberis type), chemically dominated by nonacosan-10-ol, or irregular platelets or as parallel grouped (usually non-entire) platelets (Hypericum type). Secretory cavities usually absent. Mesophyll usually with sclerenchymatous idioblasts (with asterosclereids, columellar or palisade sclereids). Idioblasts with ethereal oils absent. Leaf margin serrate (sometimes spinose-dentate), crenate or entire. Extrafloral nectaries sometimes present.

Inflorescence Terminal or axillary, with flowers solitary or pairwise (representing reduced inflorescence branches; rarely with terminal flower as well) in spike-, spadix-, cone- or head-like, or compound inflorescence. Numerous species have pseudanthium with involucre consisting of often sepaloid to petaloid bracts.

Flowers Usually zygomorphic (often somewhat obliquely; rarely actinomorphic or asymmetric). Hypanthium present (calyx tube with stamens inserted at adaxial side) or absent. Hypogyny or perigyny. Sepals four, with valvate aestivation, petaloid, free or often connate in lower part into a tube (all four sepals or 3+1). Small, almost spiny, abaxial projection,‘Vorläuferspitze’, sometimes present immediately below tepal apex. Petals probably absent; in their place (two to) four hypogynous alternisepalous scales or glands, free or connate into annular often quadrilobate extrastaminal nectariferous disc (some species without nectaries). Receptacular nectaries present.

Androecium Stamens four, haplostemonous, antesepalous. Filaments usually free, mostly entirely or partially adnate to tepals (in Bellendena free from sepals). Anthers usually free (sometimes connivent, rarely connate?), basifixed, non-versatile, usually tetrasporangiate (lateral anthers often disporangiate; in Conospermum and Synaphea all disporangiate), usually introrse (rarely latrorse), longicidal (dehiscing by longitudinal slits); connective sometimes prolonged. Tapetum secretory. Staminodia usually absent (one or several staminodia present in some species).

Pollen grains Microsporogenesis simultaneous. Pollen grains usually triporate (sometimes di- to octoporate, or di- to octocolporate; rarely di- to octocolpate; in Beauprea tricolpoidate), shed as monads, bicellular at dispersal. Pores widely operculate, arranged according to Garside’s rule (known also in Santalales): apertures produced in threes at four points in tetraedral tetrad (in majority of Triaperturate Fischer’s rule: apertures produced pairwise on six points in tetrad); diporate pollen grains (in Embothrium and Banksieae) due to change from simultaneous to successive microsporogenesis and from tetraedral to decussate or tetragonal arrangement of microspores. Exine tectate or semitectate, with columellate infratectum, reticulate or scabrate, echinate. Endexine usually absent (often present in Grevilleoideae).

Gynoecium Pistil composed of a single, entirely or partially closed carpel, usually stipitate (gynophore); carpel plicate, postgenitally entirely fused, without canal. Ovary superior or semi-inferior, monocarpellate and unilocular. Stylodium usually single (rarely absent), usually with apex expanded around stigma and specialized for secondary pollen presentation. Stigma single, terminal, subterminal or lateral (often slit-like), usually papillate, Dry or Wet type. Pistillodium absent?

Ovules Placentation marginal eller subapical (to subbasal?). Ovules usually one to three (rarely up to 100 or more) per carpel, hemianatropous, anatropous, amphitropous or orthotropous (micropyle directed downwards), pendulous, bitegmic, crassinucellar. Micropyle endostomal. Outer integument two to nine cell layers thick. Inner integument two or three cell layers thick. Vascular bundles forming chalazal ring. Nucellar cap endothelial, approx. four cell layers thick. Megagametophyte Polygonum type. Position of megagametophyte very varying. Synergids usually with a filiform apparatus. Antipodal cells sometimes persistent. Endosperm formation ab initio nuclear, later usually entirely or partially cellular. Endosperm haustorium chalazal; megasporangial cells usually gradually degenerating, and endosperm without haustorium (megasporangial cells surrounding megagametophyte in Grevilleoideae rapidly degenerating, endosperm haustoria usually developed; in Lomatia, unicellular finger-like processes formed all over endosperm surface). Embryogenesis asterad.

Fruit A follicle, capsule, achene or drupe, often fused into a cone-like syncarp (often lignified and long persistent; often dehiscing only after fire). Pericarp and integument sometimes fused forming a caryopsoid fruit.

Seeds Aril absent. Elaiosome present in some species. Seed winged or unwinged. Seed coat usually testal. Testa sometimes multiplicative, occasionally reduced. Exotesta sometimes palisade. Endotesta often palisade, with fibrillar cells and calcium oxalate crystals. Exotegmen often fibrous. Endotegmen? Suspensor absent. Perisperm not developed. Endosperm usually absent (present, although weakly developed, in at least Bellendena and Persoonia). Embryo large, straight, oily and proteinaceous (starch absent), well differentiated, without chlorophyll. Cotyledons usually two (in Persoonia and Toronia three or more, in some species of Persoonia three to nine), large. Germination phanerocotylar or cryptocotylar?

Cytology n = 5, 7, 10–14, 26, 28 – Chromosomes sometimes very large (particularly in Persoonioideae).

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), flavonol derivatives, cyanidin, condensed tannins, ellagitannins, proanthocyanidins (prodelphinidins), tropane alkaloids (bellendine present in Bellendena; methylbellendine and 2α-benzoyltropane present at least in Darlingia; benzoyltropine, benzoyltropane and phenylhydroxymethyl tropane present at least in Knightia and Darlingia), saponins, sapogenins, naphthoquinones, benzoquinones (rapanone, arbutin), tyrosine derived cyanogenic glycosides (dhurrin, proteacin, taxiphyllin, triglochinin), and quebrachitol present. Benzylisoquinoline alkaloids not found. Aluminium accumulated in many species of Grevilleoideae and Placospermum.

Use Ornamental plants, seeds (Macadamia integrifolia, M. tetraphylla), medicinal plants, dyeing substances, timber.

Systematics Proteaceae are sister-group to Platanus (Platanaceae). DNA sequence analyses of Proteaceae (Hoot & Douglas 1998) give the topology [Bellendenoideae+[Persoonioideae+[Symphionematoideae+[Proteoideae+Grevilleoideae]]]]. On the other hand, Weston & Barker (2006) identified the sister-group relationship [Bellendena+Persoonioideae].

Bellendenoideae P. H. Weston, Fl. Austral. 16: 472. 30 Nov 1995

1/1. Bellendena (1; B. montana; Tasmania). – Proteoid (cluster) roots present. Stomata laterocytic. Inflorescence a terminal raceme. Bracts absent. Filaments not adnate to tepals. Hypogynous glands absent. Ovules two per carpel. Fruit dry, winged, indehiscent. Endosperm usually present (weakly developed). Cotyledons not auriculate. n = 5. Chromosomes 6,5–7 μm long. Bellendine present. Aluminium accumulated.

[Persoonioideae+[Grevilleoideae+[Symphionematoideae+Proteoideae]]]

Stomata brachyparacytic. Tepals connate. Filaments adnate to tepals. Connective sometimes with appendage. Four nectary lobes usually present. Stylodia long. Endosperm present.

Persoonioideae Engl. in Engler et Prantl, Nat. Pflanzenfam. III, 1: 128. Mai 1888

2/110. Placospermeae C. T. White et W. D. Francis in Proc. roy. Soc. Queensland 35: 79. 1924. Placospermum (1; P. coriaceum; northeastern Queensland). – Persoonieae Endl., Gen. Plant.: 339. Dec 1837. ’Persoonia’ (c 100; paraphyletic?; Australia, New Caledonia (Garnieria) and New Zealand; incl. Toronia, Garnieria and Acidonia). – Australia, New Caledonia, New Zealand. Proteoid (cluster) roots absent (secondarily lost). Vestured pits present. Filaments entirely or largely adnate to tepals. Anthers free or basally (or entirely) adnate to tepals. Hypogynous glands present. Secondary pollen display absent. Ovules usually one or two (sometimes more than two) per carpel. Fruit usually a drupe (in Placospermum a follicle). Cotyledons obreniform. n = 7 (rarely n = 14). Mean length of chromosomes 9,1–14,4 μm. Aluminium accumulated in Placospermum.

[Grevilleoideae+[Symphionematoideae+Proteoideae]]

Flowers vertically or obliquely zygomorphic: tepals on one side split to base into four, or three tepals connate and one tepal free. Secondary pollen display present in some species (stylar apex collecting pollen grains). Ovules sometimes anatropous. x = 14. Mean length of chromosomes 0,5–5 μm.

Grevilleoideae Engl. in Engler et Prantl, Nat. Pflanzenfam. III, 1: 128. Mai 1888

45/855. Roupaleae Meisn., Plant. Vasc. Gen.: Tab. Diagn. 332, Comm. 245. 18-24 Jul 1841 [’Rhopaleae’]. Megahertzia (1; M. amplexicaulis; northeastern Queensland), Knightia (1; K. excelsa; New Zealand), Eucarpha (2; E. deplanchei, E. strobilina; New Caledonia), Triunia (4; T. erythrocarpa, T. montana, T. robusta, T. youngiana; eastern Queensland, northeastern New South Wales), Roupala (33; Mexico, Central America, Andean South America south to Argentina), Neorites (1; N. kevedianus; Queensland, southeastern New South Wales), Orites (9; eastern Queensland, eastern New South Wales, Victoria, Tasmania, Bolivia, Andean Chile), Lambertia (10; southwestern Western Australia, New South Wales), Xylomelum (7; X. angustifolium, X. benthamii, X. cunninghamianum, X. occidentale, X. pyriforme, X. salicinum, X. scottianum; western and eastern Australia), Helicia (c 100; South, East and Southeast Asia, Malesia to New Guinea, Australia), Hollandaea (4; H. diabolica, H. porphyrocarpa, H. riparia, H. sayeriana; northeastern Queensland), Darlingia (2; D. darlingiana, D. ferruginea; northern Queensland), Floydia (1; F. praealta; southeastern Queensland, northeastern New South Wales). – Banksieae Dumort., Anal. Fam. Plant.: 18. 1829. Musgravea (2; M. heterophylla, M. stenostachya; northeastern Queensland), Austromuellera (2; A. trinervia, A. valida; northeastern Queensland), Banksia (c 170; northern, southwestern and eastern Australia, Tasmania). – Embothrieae Meisn. in A. P. de Candolle et A. L. P. P. de Candolle, Prodr. 14: 211, 443. Oct (med.) 1856.Lomatia (12; eastern Australia, Tasmania, Peru, Chile, western Argentina), Embothrium (2–8; E. coccineum, E. lanceolatum; southern Peru, Chile, western Argentina), Oreocallis (1; O. grandiflora; the Andes in Ecuador and Peru), Alloxylon (4; A. brachycarpum, A. flammeum, A. pinnatum, A. wickhamii; New Guinea, Aru Islands, eastern Queensland, eastern New South Wales), Telopea (5; T. aspera, T. mongaensis, T. oreades, T. speciosissima, T. truncata; New South Wales, Victoria, Tasmania), Stenocarpus (21; New Guinea, Aru, northern and eastern Australia, New Caledonia), Strangea (3; S. cynanchicarpa, S. linearis, S. stenocarpoides; southwestern Western Australia, southeastern Queensland, northeastern New South Wales), Opisthiolepis (1; O. heterophylla; northeastern Queensland), Buckinghamia (2; B. celsissima, B. ferruginiflora; northeastern Queensland), Grevillea (c 360; Malesia to New Guinea, Australia, New Caledonia; paraphyletic?; incl. Hakea and Finschia?), Hakea (c 150; Australia; in Grevillea?), Finschia (3; F. chloroxantha, F. ferruginiflora, F. rufa; New Guinea, the Bismarck Archipelago, Solomon Islands, Aru, Palau, Vanuatu; in Grevillea?). – Macadamieae Venkata Rao in Proc. Indian Acad. Sci. 48B: 23. 1968. Macadamia (9; Madagascar, Sulawesi, Queensland, New South Wales, New Caledonia; paraphyletic?; incl. Panopsis and Brabeium?), Panopsis (c 25; Central America, tropical South America; in Macadamia?), Brabeium (1; B. stellatifolium; Western Cape; in Macadamia?), Malagasia (1; M. alticola; Madagascar), Catalepidia (1; C. heyana; northeastern Queensland), Virotia (6; V. angustifolia, V. francii, V. leptophylla, V. neurophylla, V. rousselii, V. vieillardi; New Caledonia), Athertonia (1; A. diversifolia; northeastern Queensland), Heliciopsis (14; southeastern China, Burma, Thailand, Indochina, Malesia to the Philippines), Cardwellia (1; C. sublimis; northeastern Queensland), Sleumerodendron (1; S. austrocaledonicum; New Caledonia), Euplassa (c 20; South America, especially northwestern Andes, the Guayana Highlands and southeastern Brazil), Gevuina (1; G. avellana; southern Chile, southwestern Argentina), Bleasdalea (5; B. bleasdalei, B. ferruginea, B. lutea, B. papuana, B. vitiensis; New Guinea, northeastern Queensland), Hicksbeachia (2; H. pilosa, H. pinnatifolia; southeastern Queensland, northeastern New South Wales), Kermadecia (4; K. elliptica, K. pronyensis, K. rotundifolia, K. sinuata; New Caledonia). – Unplaced Grevilleoideae Sphalmium (1; S. racemosum; northeastern Queensland), Carnarvonia (1; C. araliifolia; northeastern Queensland). – South Africa, Madagascar, southern India, Sri Lanka, Southeast Asia to New Guinea, Australia, Tasmania, Melanesia, Mexico to South America. Proteoid (cluster) roots present. Sieve elements with rosette-shaped non-dispersive protein bodies. Inflorescence usually a raceme of paired flowers (often subtende by a common bract) or a panicle of such racemes. Filaments basally to entirely adnate to tepals. Pollen grains with endexine often present also in apertural regions. Ovules (one or) two or several per carpel. Fruit a non-winged follicle with winged seeds, or a drupe. Seeds sometimes pachychalazal. Endosperm with chalazal nuclear haustorium. Cotyledons usually auriculate at base (rarely peltate). n = (10–)14(–15). Chromosomes 1,0–2,6 μm long. Aluminium accumulated in many species (not in Embothrieae). – Carnarvonia (Carnarvonioideae L. A. S. Johnson et B. G. Briggs in Bot. J. Lionn. Soc. 70: 172. 3 Sep 1975) and Sphalmium (Sphalmioideae L. A. S. Johnson et B. G. Briggs in Bot. J. Linn. Soc. 70: 172. 3 Sep 1975) are provisionally included in Grevilleoideae, yet morphologically very distinct and possibly sister-groups.

[Symphionematoideae+Proteoideae]

Fruit indehiscent, dry.

Symphionematoideae P. H. Weston et N. P. Barker in Telopea 11: 330. 13 Nov 2006

2/3. Agastachys (1; A. odorata; western Tasmania), Symphionema (2; S. montanum, S. paludosum; eastern New South Wales). – Southeastern Australia, Tasmania. Proteoid (cluster) roots absent. Spicate inflorescence (in Symphionema often compound). Hypogynous glands (nectaries) absent. Filaments adnate at base to tepals. Ovules one or two per carpel. Cotyledons not auriculate. n = 10, 14. Chromosomes c. 3 μm long.

Proteoideae Eaton, Bot. Dict., ed. 4: 30. Apr-Mai 1836 [‘Proteae’]

25/640. Conospermeae Endl., Gen. Plant.: 338. Dec 1837. Stirlingia (7; S. abrotanoides, S. anethifolia, S. divaricatissima, S. latifolia, S. seselifolia, S. simplex, S. tenuifolia; southwestern Western Australia), Conospermum (53; Australia), Synaphea (c 50; southwestern Western Australia). – Petrophileae P. H. Weston et N. P. Barker in Telopea 11: 331. 13 Nov 2006.Petrophile (53; southwestern, southern and eastern Australia), Aulax (3; A. cancellata, A. pallasia, A. umbellata; Western Cape). – Proteeae Dumort., Anal. Fam. Plant.: 18. 1829 [‘Proteae’]. Protea (103 or 114; eastern and southern Africa), Faurea (c 15; Africa, Madagascar). – Leucadendreae P. H. Weston et N. P. Barker in Telopea 11: 332. 13 Nov 2006. Isopogon (c 35; Australia), Adenanthos (33; southwestern Western Australia, South Australia, western Victoria), Leucadendron (80–85; Western and Eastern Cape, KwaZulu-Natal), Serruria (c 50; Western Cape), Paranomus (18–19; Western Cape, southern Eastern Cape), Vexatorella (4; V. alpina, V. amoena, V. latebrosa, V. obtusata; southern Northern Cape, central Western Cape), Sorocephalus (11; Western Cape), Spatalla (20; Western Cape), Leucospermum (c 50; Western and Eastern Cape, KwaZulu-Natal, Zimbabwe), Mimetes (13; Western Cape), Diastella (7; D. buekii, D. divaricata, D. fraterna, D. myrtifolia, D. parilis, D. proteoides, D. thymelaeoides; Western Cape), Orothamnus (1; O. zeyheri; the Kogelberg area in Western Cape); incertae sedis: Eidothea (2; E. hardeniana, E. zoexylocarya; northeastern Queensland, northeastern New South Wales), Beauprea (13; New Caledonia), Beaupreopsis (1; B. paniculata; New Caledonia), Dilobeia (2; D. tenuinervis, D. thouarsii; Madagascar), Cenarrhenes (1; C. nitida; Tasmania), Franklandia (2; F. fucifolia, F. triaristata; southwestern Western Australia). – Africa south of Sahara (with their highest diversity in the Cape region), Madagascar, Australia. Sometimes herbaceous. Proteoid (cluster) roots present. Sieve elements with non-dispersive protein bodies. Inflorescence basically racemose, often compound, often condensed into spikes or capitula. Flowers sessile. Hypanthium sometimes present. Filaments poorly to entirely adnate to tepals. Anthers sometimes monothecal. Ovule usually one (in some species of Petrophile and Isopogon sometimes two) per carpel. Fruit often a one-seeded drupe or nut. Cotyledons not auriculate. n = (10–)11–13(–14). Chromosomes 1,2–3,4 μm long. Aluminium rarely accumulated.

Cladogram (simplified) of Proteaceae based on DNA sequence data (Hoot & Douglas 1998). Bellendena is sister to Persoonioideae in analyses by, e.g., Weston & Barker (2006).

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PROTEALES Juss. ex Bercht. et J. Presl

NELUMBONACEAE A. Rich.

PLATANACEAE T. Lestib.

PROTEACEAE Juss.