ERICALES Dumortier Main Tree, Synapomorphies

Ericopsida Bartl., Ord. Nat. Plant.: 120, 152. Sep 1830 [’Ericineae’]; Ericanae Takht., Sist. Filog. Cvetk. Rast. [Syst. Phylog. Magnolioph.]: 221. 4 Feb 1967; Theanae R. Dahlgren ex Reveal in Phytologia 74: 179. 25 Mar 1993; Primulanae R. Dahlgren ex Reveal in Phytologia 79: 71. 29 Apr 1996; Theidae Doweld, Tent. Syst. Plant. Vasc.: xliii. 23 Dec 2001; Ericidae C. Y. Wu in Acta Phytotaxon. Sin. 40: 308. 2002

Fossils Actinocalyx bohrii from the Late Santonian to the Early Campanian of Sweden comprises hypogynous flowers with a pentamerous biserial perianth, five haplostemonous stamens and three connate carpels. The five petals are basally connate and the filaments are adnate to the corolla. The pollen grains are tricolpate with a finely foveolate exine. The ovary is trilocular, the placentation is central, the ovules are anatropous and the fruits are capsular. Paradinandra suecica from the Late Santonian to the Early Campanian of Sweden resembles Actinocalyx, although it has basally connate stamens which are arranged in an outer whorl of five and an inner whorl of ten stamens. The paired stamens were probably antepetalous and solitary stamens were possibly likewise antepetalous. There is an intrastaminal nectariferous disc. The anthers probably dehisced by apical pores or slits, and the pollen grains were tricolpate. The placentation is intrusively parietal and the ovules are campylotropous.

Paleoenkianthus sayrevillensis, from the Turonian of New Jersey, is represented by hypogynous pentamerous flowers and fruits. The petals are basally connate and have a valvate aestivation. There is one series of eight stamens with inverted anthers bearing two or three appendages at their base, and the pollen grains are probably tricolporate with reticulate exine. The four connate carpels form a quadrilocular ovary with four free styles, and developed into a capsule with septicidal-loculicidal dehiscence.

Pentapetalum trifasciculandricum, from the Turonian of New Jersey, includes pentamerous hypogynous flowers with a large number of stamens in three fascicles, and three carpels forming free styles and a trilocular ovary containing numerous ovules. ‘Pentaphylax’ protogaea, fossil fruits from the Maastrichtian of Germany, has been assigned to the extant Pentaphylacaceae, although there are several important differences. The fossils have a thick lignified pericarp and the persistent calyx lobes have unequal length. Moreover, the seeds are anatropous.

Habit Bisexual, monoecious, polygamomonoecious, dioecious, polygamodioecious or functionally monoecious, dioecious or gynodioecious, evergreen or deciduous trees, shrubs, suffrutices or lianas, or perennial, biennial or annual herbs.

Vegetative anatomy Usually with arbuscular mycorrhiza with Glomales. Phellogen ab initio superficially or deeply seated. Secondary lateral growth normal or absent. Vessel elements with scalariform or simple (sometimes reticulate) perforation plates; lateral pits usually alternate, scalariform or opposite, simple or bordered pits (vessels occasionally absent). Pit membrane remnants may occur in some species. Imperforate tracheary xylem elements tracheids or fibre tracheids with bordered pits, non-septate (sometimes also vasicentric tracheids). Wood rays uniseriate or multiseriate, usually heterocellular (sometimes homocellular), or absent. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty vasicentric, reticulate, unilateral or banded (rarely aliform, confluent or scalariform), or absent. Sieve tube plastids usually Ss type (sometimes Pcf type). Nodes usually 1:1, unilacunar with one trace (rarely 3:3, trilacunar with three traces). Idioblasts with mucilage often present. Laticifers with milky latex occasionally present; ducts with gutta-percha sometimes present. Parenchyma often with idioblasts containing calciumoxalate raphides. Prismatic or acicular crystals, crystal sand, styloids or druses sometimes present.

Trichomes Hairs unicellular or multicellular, uniseriate or multiseriate, simple or branched, bristle-like, flattened, furcate, stellate or dendritic (sometimes fasciculate, T-shaped, candelabra-, funnel- or cup-shaped, rarely arachnoid, peltate or lepidote), or absent; multicellular glandular hairs with capitate head (sometimes thick and colleter-like, rarely peltate-lepidote) sometimes present.

Leaves Usually alternate (spiral or sometimes distichous, rarely tetrastichous; rarely opposite or verticillate), simple, entire, sometimes coriaceous or ericoid (rarely scale-like), with conduplicate, supervolute, involute, convolute or revolute (rarely circinnate or plicate) ptyxis. Stipules very small and caducous or absent; leaf sheath absent. Colleters often abundant. Petiole vascular bundle transection arcuate or annular (sometimes D-shaped or complex, sometimes with one or two medullary bundles). Venation pinnate, eucamptodromous, brochidodromous, craspedodromous or semicraspedodromous (rarely reticulodromous, acrodromous to palinactinodromous or actinodromous), or leaves single-veined; vascular bundles often collateral; some veins proceeding into teeth. Stomata usually anomocytic (sometimes paracytic, anisocytic, cyclocytic or actinocytic, rarely stephanocytic, staurocytic or parallelocytic). Cuticular wax crystalloids as platelets or rodlets (sometimes amorphous or as tubuli dominated by diketones), or absent. Epidermis with or without mucilage cells. Mesophyll often with calciumoxalate raphides. Mesophyll with or without sclerenchymatous idioblasts containing brachysclereids, osteosclereids and other kinds of sclereids. Secretory oil cavities present or absent. Ducts with gutta-percha sometimes present. Leaf margin usually serrate or entire (sometimes crenate). Leaf tooth apex often widened, translucent, persistent, sometimes gland-tipped (rarely with nectariferous glands); teeth often theoid, with single vein and opaque deciduous cap (sometimes with single multicellular hair replacing cap).

Inflorescence Usually axillary (sometimes terminal), panicle, thyrsopaniculate, thyrsoid, botryoid, fasciculate, corymbose, head-, umbel-, raceme- or spike-like, cymose or racemose (flowers sometimes solitary, axillary or terminal). Floral prophylls (bracteoles) present or absent.

Flowers Actinomorphic or zygomorphic. Usually hypogyny (sometimes epigyny or half epigyny). Sepals (two to) five (to twelve), with imbricate or valvate (sometimes open, rarely contorted) aestivation, usually persistent, free or more or less connate. Petals (three to) five (to 18), with imbricate, valvate or contorted (rarely conduplicate or subinduplicate) aestivation, usually caducous, free or more or less connate into hypocrateromorphous, discoid, campanulate, infundibuliform, tubular or urceolate corolla (sometimes nectariferous at base). Nectary and disc usually absent (nectariferous disc sometimes annular or consisting of separate parts, intrastaminal or occasionally extrastaminal, or petals sometimes nectariferous).

Androecium Stamens (two to) five to more than 1.200, in one or more whorls (sometimes in fascicles). Filaments usually free (rarely connate at base), usually free from tepals (sometimes epipetalous). Anthers dorsifixed, ventrifixed or subbasifixed, versatile or non-versatile, often inverting and finally resupinate, usually tetrasporangiate (rarely disporangiate), extrorse or introrse (rarely latrorse), longicidal (dehiscing by longitudinal or transverse slits) or poricidal (dehiscing by seemingly apical to subapical pores). Tapetum usually secretory (rarely amoeboid-periplasmodial). Staminodia sometimes present.

Pollen grains Microsporogenesis simultaneous. Pollen grains (2–)3(–6)-colpor(oid)ate, (2–)3(–5)-colpate or 3(–5)-porate (rarely pantocolporate, pantoporate or stephanocolpate), usually shed as monads (sometimes tetrads, rarely dyads, triads or polyads), usually bicellular (rarely tricellular) at dispersal. Exine tectate or semitectate (rarely intectate), usually with granular (sometimes columellate) infratectum, pertectate, reticulate, foveolate, scabrate, fossulate, areolate, rugulate, psilate, gemmate, verrucate, verruculate, echinate, scrobiculate, granulate or microgranulate.

Gynoecium Pistil composed of (one to) five to c. 20 (to c. 30) more or less connate carpels. Ovary usually superior (sometimes inferior or semi-inferior), (unilocular to) quinquelocular to c. 20(–30)-locular (sometimes incompletely septate). Style single, simple, or stylodia (three to) five to c. 20, short, free or partially connate, sometimes persistent. Stigma capitate, punctate, peltate or lobate, papillate or non-papillate, Dry or Wet type. Male flowers sometimes with pistillodium.

Ovules Placentation usually axile (sometimes apical or parietal, rarely basal; placentae often diffuse). Ovules one to numerous per carpel, usually anatropous (sometimes hemianatropous or campylotropous, rarely orthotropous), pendulous to ascending, apotropous or epitropous, usually unitegmic (sometimes bitegmic), tenuinucellar. Micropyle endostomal or bistomal (when bitegmic usually endostomal). Megagametophyte usually monosporous, Polygonum type (sometimes disporous, Allium type, rarely tetrasporous, Adoxa type). Synergids sometimes with a filiform apparatus. Antipodal cells sometimes persistent and proliferating. Endosperm development usually cellular (sometimes nuclear). Endosperm haustoria chalazal and/or micropylar or absent. Embryogenesis solanad, onagrad or asterad (sometimes caryophyllad or chenopodiad).

Fruit A loculicidal and/or septifragal capsule (rarely a pyxidium or a denticidal capsule), a berry, a drupe or a samara (rarely a nut enclosed by perianth or a drupaceous schizocarp), often with persistent and sometimes accrescent calyx.

Seeds Aril usually absent. Exotesta with inner cell walls of outer epidermis often thickened, with theoid exotestal thickenings. Perisperm not developed. Endosperm copious to sparse (rarely ruminate), oily or proteinaceous. Embryo large to small, usually straight (sometimes curved), well differentiated, usually without chlorophyll. Cotyledons two. Germination usually phanerocotylar (rarely cryptocotylar).

DNA Nuclear gene PI duplicated. I copy of nuclear gene RPB2 present. Mitochondrial intron coxII.i3 often lost.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin, gossypetin), catechins, cyanidin, davidigenin, Route I secoiridoids, Group I carbocyclic iridoids (aucubin, daphylloside, monotropein), Group II decarboxylated iridoids (e.g. cornine), Group IV carbocyclic iridoids (rarely unedoside, a C8 iridoid glycoside), Group VI secoiridoids (morroniside), Group X secoiridoids (nepetalactones, iridoidpyridine alkaloids), sarracenin, oleanolic acid derivatives, monoterpenes, diterpenes (e.g. andromedotoxin), sesquiterpenes, triterpenes, free triterpenic acids (betulinic acid, ursolic acid), dammaranes, arjunolic acid derivatives, ellagic acid (frequent to absent), gallic acid, ellagitannins (geraniin, pedunculagin, tellimagrandin II), non-hydrolyzable tannins, proanthocyanidins (prodelphinidins), caffeic acid, chlorogenic acid, phenolic heterosides, 3-galactoside, triterpene saponins, benzoquinones (e.g. arbutin, rapanone) and naphthoquinones and their derivatives (e.g. naphthoquinone derivatives of 7-methyljugone, lawsone, plumbagin, droserone), anthraquinones, polyacetate-derived arthroquinones, benzopyrones, cucurbitacins, dihydrosterculic acid, lignans (pinoresinol), myo-inositol, and ethereal oils present. Alkaloids (e.g. pyrrolizidine alkaloids, indole alkaloids and their glycosides) and cyanogenic compounds rare. Carbohydrates sometimes stored as oligosaccharides with kestose or isokestose bonds (inulin). Aluminium accumulated in some groups.

The clade [Balsaminaceae+[Marcgraviaceae+Tetrameristaceae]] is sister-group to all other Ericales. Potential synapomorphies are, according to Stevens (2001 onwards): vessel elements with usually simple perforation plates; vessels in radial multiples; presence of raphid sacs in stem; presence of branched sclereids; absence of druses; leaf with supervolute ptyxis; lamina elongating in bud and with obscure abaxial lines; leaf margin serrate; inflorescence racemose; abaxial petal surface with stomata; filiform appendages present along stomium; stamens as many as sepals, antesepalous, free from petals; anthers subbasifixed; ovary with mucilaginous secretions; absence of gynoecial nectary; style short, stigma little expanded; ovules bitegmic; micropyle endostomal; presence of micropylar endosperm haustorium; presence of myricetin; and absence of ellagic acid. Balsaminaceae and Tetrameristaceae share the synapomorphies: sepals petaloid, nectariferous; filaments postgenitally connate; and stylar canal present, quinqueradiate.

The leaves in this basal clade are often glabrous, simple and usually lack stipules (some Balsaminaceae have stipules). Extrafloral nectaries or glands are often present on the adaxial side of leaf (including petiole), calyx or corolla. The pollen grains are tricolporate and bicellular at dispersal, and the ovules are bitegmic and tenuinucellar. The endosperm development is cellular, endosperm haustorium is present, and the endosperm is sparse. The nodes are unilacunar in Balsaminaceae and Marcgraviaceae, and there is a caducous calyptra, developed from androecium (Balsaminaceae) or petals (Marcgraviaceae). Marcgraviaceae and Tetrameristaceae have indehiscent fruit, and Pelliciera and Marcgraviaceae have large cortical air cavities.

The Ericales main clade may be characterized by connate corolla with well developed tube, and long style. The clade [Polemoniaceae+Fouquieriaceae] is part of a trichotomy forming the base of the remaining Ericales. Fouquieria and Polemoniaceae share the potential synapomorphies (Stevens 2001 onwards): phellogen outer-cortical; vessel elements with simple perforation plates; inflorescence terminal, cymose; hypogyny; sepals free, with membranous margins and abaxial stomata; petals largely connate; presence of nectaries; filaments adnate to corolla; anther thecae largely separate, without septum; pistil composed of three connate carpels; presence of gynoecial nectary; style hollow, strongly lobate; ovules arranged in two ranks, apotropous; fruit a loculicidal capsule, with persistent calyx and seeds inserted on central columella; seed coat winged; endosperm sparse; and loss of mitochondrial intron coxII.i3. Lecythidaceae are the second lineage of the trichotomy, whereas the third monophyletic group comprises the main clade of Ericales.

The first of three clades in a basal trichotomy forming the remainder of Ericales has the topology [Sladeniaceae+[Pentaphylacaceae+Ternstroemiaceae]]. This clade has the following potential synapomorphies (Stevens 2001 onwards): vessel elements with scalariform perforation plates; bordered vessel-fibre pits; nodes 1:1; presence of mucilage cells; unicellular hairs; petiole vascular bundle transection arcuate; campanulate corolla; petals less than one centimetre long and connate only at base; absence of nectary; basifixed anthers; pollen grains 14–30 μm long; exine usually only little ornamented; placentae becoming swollen; bitegmic ovules; endostomal micropyle; inner integument three or four cell layers thick; fruit a capsule with columella and calyx persistent; presence of endosperm; and long embryo. Pentaphylax and Ternstroemiaceae share the following synapomorphies: parenchyma apotracheal; intervessel pitting opposite-scalariform; leaves with supervolute ptyxis; flowers solitary axillary or inflorescence fasciculate; corolla usually greenish to yellowish; style hollow; ovules campylotropous to hemitropous; mesotesta well developed; embryo U-shaped; and accumulation of aluminium.

The second clade in this trichotomy has the following topology: [Sapotaceae+ [Ebenaceae+[Maesaceae+[[Theophrastaceae+Samolaceae]+[Myrsinaceae+Primulaceae]]]]]. Morphological and phytochemical characters supporting this clade are few: connate petals; apotropous ovules; and absence of ellagic acid. Furthermore, bitegmic ovules and inner integument thicker than outer are two potential synapomorphies uniting Ebenaceae with the following clade. [Maesaceae+[[Theophrastaceae+Samolaceae]+[Myrsinaceae+Primulaceae]]] is a very strongly supported clade. Synapomorphies are, e.g., the following (Stevens 2001 onwards): schizogenous secretory canals with yellow, red or brown substances (e.g. tannins) sometimes present; nodes 3:3?; stomata sometimes anisocytic; presence of small immersed and often peltate glandular hairs; inflorescence racemose; petals and stamens arising from common primordia; presence of nectaries; stamens as many as petals, antepetalous; staminodia antesepalous, represented by at least a vascular trace; carpels five, antepetalous; style short, hollow; stigma capitate; placentation free central, with large placenta; ovules at least partially immersed in swollen placenta, apotropous, bitegmic; micropyle bistomal; endosperm development nuclear; tanniniferous endothelium present; seeds angular; endotesta crystalliferous; endosperm copious, sometimes with amyloid (xyloglucans) or hemicellulose, and sometimes with thick and pitted cell walls.

The clade [[Theophrastaceae+Samolaceae]+[Myrsinaceae+Primulaceae]] is characterized by wood rays at least 5-seriate; absence of uniseriate rays in woody species; presence of small immersed often peltate or glandular hairs; absence of floral prophylls (bracteoles); corolla subrotate, with imbricate aestivation; corolla tube fairly short; petals arising abaxially from common primordium; and staminal primordium larger than petal primordium. Theophrastaceae and Samolus share the characters bracts displaced up pedicels and presence of petaloid staminodia, whereas Myrsinaceae och Primulaceae share two identical deletions in plastid genendhF.

The third clade of the trichotomy comprises (1) Theaceae, (2) the clade [Symplocaceae+ [Styracaceae+Diapensiaceae]] and (3) the clade [Sarraceniaceae+[Actinidiaceae+Roridulaceae]]+ [Clethraceae+[Cyrillaceae+Ericaceae]] and may be characterized by the synapomorphies: vessel elements with scalariform perforation plates, and serrate leaves. An additional lineage is the mycotrophical holoparasitic Mitrastemma although with a highly uncertain position.

The clade [Symplocaceae+[Styracaceae+Diapensiaceae]] is characterized by the features: woody habit; racemose inflorescence; hollow style; and copious endosperm. The lineage [Styracaceae+Diapensiaceae] has pericyclic phellogen; nodes 1:1; absence of glandular hairs; spiral leaves with usually serrate margin; nectariferous disc absent or poorly developed and annular; stamens usually twice as many as petals; filaments connate at base or adnate to corolla tube; anthers basifixed, introrse, longicidal; endothecium well developed and with fibrous thickenings; pollen grains tricolporate, shed as monads and usually bicellular at dispersal; pistil composed of two to four carpels; hollow style continuing without interruption into locules; ovules numerous per carpel; unitegmic, tenuinucellar; megagametophyte usually with large amount of starch; endosperm development cellular; and fruit usually a loculicidal capsule. Furthermore, calciumoxalate crystals are often present as aggregations in idioblasts in the parenchyma.

The [[Sarraceniaceae+[Actinidiaceae+Roridulaceae]]+[Clethraceae+[Cyrillaceae+Ericaceae]]] clade has the potential synapomorphies (Stevens 2001 onwards): inflorescence racemose; anthers extrorse, inverting during development, and dehiscing by apical pores or short slits; pollen grains rugulate; tectum and foot layer solid; infratectum granular; pistil composed of three carpels, with median carpel adaxial, or five antepetalous carpels; style impressed into ovary apex; ovules numerous per carpel; endothelium present; fruit a capsule; exotesta with strongly thickened inner cell walls; endosperm copious; and absence of mitochondrial intron coxII.i3. The clade [Sarraceniaceae+[Actinidiaceae+Roridulaceae]] is characterized by the features: absence of nectary; stigma Dry type; presence of hypostase; and presence of Route I secoiridoids. The lineage [Clethraceae+[Cyrillaceae+Ericaceae]] has the synapomorphies: phellogen pericyclic; absence of pericyclic fibres; leaves spiral; inflorescence racemose; absence of floral prophylls; flowers pendulous; basal part of ovary wall nectariferous; stamens twice as many as sepals; style hollow; endosperm with micropylar and chalazal haustoria; testa reduced; embryo terete; and presence of ellagic acid. Finally, Cyrillaceae and Ericaceae share the synapomorphies: presence of colleters; connate petals; stigma Wet type; and presence of myricetin.

The strongly supported clade [[Sarraceniaceae+[Actinidiaceae+Roridulaceae]]+[Clethraceae+[Cyrillaceae+Ericaceae]]] has the potential synapomorphies (Stevens 2001 onwards; Löfstrand & Schönenberger 2015): inflorescence racemose; anthers attached adaxially, extrorse, inverting during development, and dehiscing by apical pores or short slits; pollen grains rugulate; tectum and foot layer solid; infratectum granular; depression at ovary-to-style transition; pistil composed of three carpels, with median carpel adaxial, or five antepetalous carpels; style impressed into ovary apex; ovules numerous per carpel; endothelium present; fruit a capsule; exotesta with strongly thickened inner cell walls; endosperm copious; and absence of mitochondrial intron coxII.i3. The clade [Sarraceniaceae+[Actinidiaceae+Roridulaceae]] is characterized by features such as: petals thick to massive; nectary absent; polystemony; stigma Dry type; hypostase present; and Route I secoiridoids present. The lineage [Actinidiaceae+Roridulaceae] is characterized by, e.g.: presence of calcium oxalate raphides; mucilage cells; inner gynoecium surface secretory; and synlateral vasculature absent from ovary. The clade [Clethraceae+[Cyrillaceae+Ericaceae]] has the synapomorphies: phellogen pericyclic; absence of pericyclic fibres; leaves spiral; inflorescence racemose; absence of floral prophylls; flowers pendulous; androecium two-worled; basal part of ovary wall nectariferous; stamens twice as many as sepals; style hollow; endosperm with micropylar and chalazal haustoria; testa reduced; embryo terete; and presence of ellagic acid. Finally, Cyrillaceae and Ericaceae share the synapomorphies: presence of colleters; connate petals; stigma Wet type; and presence of myricetin.

Gilg et Werdermann in Engler et Prantl, Nat. Pflanzenfam., ed. 2, 21: 36. Dec 1925, nom. cons.

Saurauiaceae Griseb., Grundr. Syst. Bot.: 98. 1-2 Jun 1854 [’Sauraujeae’], nom. cons.; Actinidiales Takht. ex Reveal in Phytologia 74: 174. 25 Mar 1993

Distribution Tropical and subtropical regions of East and Southeast Asia, the Himalayas, Malesia to New Guinea, northeastern Queensland, Solomon Islands, Fiji, Mexico, Central America, the Andes to Chile.

Fossils Late Cretaceous records of Actinidiaceae are known from North America and Europe and they are found at many Cenozoic sites in the Northern Hemisphere. Saurauia alenae and S. antiqua, anatropous seeds with reticulate surface, have been found in the Late Turonian to the Maastrichtian of Central Europe, and fossils similar to Actinidia are recorded from the Late Eocene onwards. Actinidiophyllum has been described from Cenozoic strata in Japan. Parasaurauia allonensis and Glandulocalyx, represented by flowers with free pentamerous perianth, ten stamens in two series and three carpels with free styles, was described from the Late Cretaceous (Late Santonian/Early Campanian) of Georgia in the United States as sister-group to [Actinidia+Saurauia] (Keller & al. 1996). Leaf impressions resembling Saurauia are known from mid-Eocene layers of North America.

Habit Bisexual, morphologically dioecious (Clematoclethra), functionally dioecious or monoecious, evergreen or deciduous trees, shrubs or lianas.

Vegetative anatomy Phellogen ab initio superficially or deeply seated. Medulla often septate by diaphragms. Endodermis prominent. Pericycle with continuous ring of sclerenchyma. Vessel elements dimorphic, with scalariform or simple (sometimes reticulate) perforation plates; lateral pits usually opposite (sometimes scalariform or alternate), simple or bordered pits. Imperforate tracheary xylem elements tracheids or fibre tracheids with bordered pits, non-septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty vasicentric or banded. Sieve tube plastids S type. Nodes usually 1:1, unilacunar with one leaf trace (rarely 3:3, trilacunar with three traces). Parenchyma with idioblasts (raphide cells) containing raphides; calciumoxalate as crystal sand present in at least Clematoclethra.

Trichomes Hairs multicellular, uniseriate, often bristle-like or flattened, or branched, stellate or dendritic; glandular hairs, short and long, sometimes present.

Leaves Usually alternate (spiral; rarely opposite), simple, entire, with conduplicate ptyxis. Stipules very small and caducous or absent; leaf sheath absent. Petiole vascular bundle transection deeply arcuate with wing bundles (Actinidia), or annular (sometimes with medullary bundles). Venation usually pinnate, camptodromous (sometimes subpalmate); some veins proceeding into teeth. Stomata usually anomocytic (sometimes paracytic). Cuticular wax crystalloids as platelets or transversely ridged rodlets. Mesophyll with calciumoxalate raphides. Leaf margin usually serrate (sometimes entire); tooth apices widened, translucent, persistent.

Inflorescence Axillary, usually thyrsopaniculate etc. (flowers sometimes solitary, axillary). Floral prophylls (bracteoles) present or absent.

Flowers Actinomorphic. Hypogyny. Sepals (three to) five (to eight), with imbricate quincuncial aestivation, usually persistent, free. Petals (three to) five (to nine), with imbricate quincuncial aestivation, caducous, free or connate at base, sometimes nectariferous at base. Nectary usually absent. Disc absent.

Androecium Stamens c. 20 to c. 240 (in Clematoclethra ten), in two or more whorls (primarily diplostemonous), usually inflexed in bud (not in Saurauia), sometimes in five antepetalous fascicles. Filaments usually free (rarely connate at base), usually free from petals (sometimes epipetalous). Anthers dorsifixed, versatile, late inverted (finally resupinate), tetrasporangiate, extrorse, longicidal (dehiscing by longitudinal or short slits widening at apex) or poricidal (dehiscing by seemingly subapical pores). Tapetum secretory, with multinucleate cells (sometimes with fused nuclei). Female flowers sometimes with staminodia?

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolporate (sometimes tetracolporate), usually shed as monads (rarely tetrads), bicellular at dispersal. Exine tectate, with granular infratectum, imperforate, psilate, microgranulate or rugulate, often transversely striate.

Gynoecium Pistil composed of (three to) five to c. 20 incompletely connate synascidiate carpels. Ovary superior, (trilocular to) quinquelocular to 20-locular (sometimes incompletely septate). Stylodia (three to) five to c. 20, short, free or partially connate, sometimes impressed, sometimes persistent, or style single, simple and hollow (Clematoclethra; in Actinidia furrowed). Stigmas capitate, peltate or lobate, papillate, Dry type. Male flowers sometimes with pistillodia?

Ovules Placentation axile. Ovules usually numerous (in Clematoclethra approx. ten) per carpel, anatropous, unitegmic, tenuinucellar. Integument six to nine cell layers thick. Hypostase well developed. Parietal tissue approx. three cell layers thick. Nucellar cap approx. three cell layers thick. Megasporocytes usually one (occasionally several, archespore multicellular). Megagametophyte monosporous, Polygonum type. Endosperm development cellular. Endosperm haustoria? Embryogenesis solanad. Polyembryony often occurring.

Fruit Usually a berry (sometimes a loculicidal capsule), often with persistent calyx; seeds embedded in pulp formed by placentae.

Seeds Aril usually absent (present in Actinidia). Testa multiplicative. Exotesta with inner cell walls of outer epidermis thick, with theoid exotestal thickenings. Endotesta? Perisperm not developed. Endosperm copious, oily or proteinaceous. Embryo large, usually straight (sometimes curved), well differentiated, without chlorophyll. Cotyledons two. Germination phanerocotylar.

Cytology n = 20, 24, 29, 30, 39 (x = 12 in Clematoclethra; x = 13 in Saurauia; x = 29 in Actinidia) – Polyploidy frequently occurring.

DNA Horizontal transfer of mitochondrial gene rps2 in Actinidia. Mitochondrial intron coxII.i3 lost.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin?), catechins, Route I secoiridoids, Group X secoiridoids (nepetalactones, iridoidpyridine alkaloids), gallic acid, proanthocyanidins (cyanidin, prodelphinidin), and polyacetate-derived arthroquinones present. Actinidin (a proteinase) present in Actinidia. Alkaloids usually absent (matatabi lactone and actinidine present). Ellagic acid, saponins, and cyanogenic compounds not found.

Richard in J. B. G. M. Bory de Saint-Vincent, Dict. Class. Hist. Nat. 2: 173. 31 Dec 1822 [’Balsamineae’], nom. cons.

Balsaminales Bercht. et J. Presl, Přir. Rostlin: 221. Jan-Apr 1820 [‘Balsamineae’]; Hydroceraceae Wilbrand, Nat. Pflanzenfam.: 26. Mai-Dec 1834 [’Hydrocereae’], nom. illeg.; Impatientaceae Lem., Ill. Hort. 1: ad t. 9. Mar 1854 [’Impatientiaceae’]; Balsaminineae Engl., Syllabus, ed. 2: 146. Mai 1898; Balsaminanae Doweld, Tent. Syst. Plant. Vasc.: xliv. 23 Dec 2001

Distribution Tropical and subtropical Africa, Madagascar, tropical Asia to New Guinea, with their largest diversity in Madagascar and Southeast Asia, few species in temperate parts of Eurasia, Africa and North America.

Habit Bisexual, although usually functionally monoecious due to extreme protandry, usually perennial or annual herbs (rarely partially woody). Stem more or less succulent, often almost translucent. Impatiens tuberosa and the epiphytic I. etindensis with large stem tubers. Numerous species are helophytes.

Vegetative anatomy Phellogen? Primary vascular tissue a cylinder of vascular bundles. Secondary lateral growth usually absent. Wood paedomorphic. Vessels solitary. Vessel elements with simple perforation plates, lateral pits? Imperforate tracheary xylem elements? Wood rays absent. Axial parenchyma? Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace. Sclerenchyma and sclereids absent. Idioblasts with mucilage present. Cortex with calciumoxalate raphides in raphid sacs.

Leaves Usually alternate (spiral; rarely opposite or verticillate), simple, entire, with involute ptyxis. Stipules and leaf sheath absent. Petiole often with glands (rarely with extrafloral nectaries or foliaceous outgrowths at leaf base, petiole or stem). Petiole vascular bundle transection arcuate. Venation pinnate. Stomata usually anomocytic (sometimes almost anisocytic). Cuticular wax crystalloids? Hydathodes abundant. Mesophyll with calciumoxalate raphides. Leaf margin usually serrate (sometimes crenate or entire); teeth apiculate, basal teeth often with nectariferous glands.

Inflorescence Axillary, raceme or umbel-like. Floral prophylls (bracteoles) sometimes absent.

Flowers Zygomorphic, usually resupinate (inverting during growth). Hypogyny. Sepals usually three (sometimes five), with imbricate aestivation, caducous, free, two lateral abaxial sepals reduced and small or absent, median adaxial sepal petaloid, usually with nectariferous spur (absent in some Malagasy species) with secretory cells on inner side, two abaxial/upper sepals very small or absent. Petals five, unequal in size, with imbricate aestivation, dorsal petal free, flat or cucullate, four lower/lateral petals usually pairwise connate (in Hydrocera free); when three sepals, then adaxial petal often with sepaloid keel. Disc absent.

Androecium Stamens five, haplostemonous, antesepalous, alternipetalous. Filaments short, stout, connate at apex and forming ring around gynoecium, upper filaments slightly longer than lower ones, free from tepals. Anthers ab initio connate into cucullate calyptra above stigma, finally coming loose at base and uplifted by accrescent gynoecium, dorsifixed to subbasifixed, non-versatile, usually dithecal (sometimes polythecal and septate by transverse trabeculae), tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains 3(–5)-colpate or 3(–5)-porate, starchy, shed as monads, usually bicellular (rarely tricellular) at dispersal. Exine semitectate, with granular infratectum, reticulate; with raphides and with cellulose threads adhering pollen grains to anther.

Gynoecium Pistil composed of (four or) five connate antepetalous carpels; when carpels four, then adaxial carpel larger. Ovary superior, (quadrilocular or) quinquelocular. Style single, very short, or absent. Stigmas usually one or (four or) five, relatively wide, non-papillate, Wet type. Pistillodium absent.

Ovules Placentation axile. Ovules usually numerous (in Hydrocera one to three) per carpel, usually anatropous (rarely hemitropous), pendulous, apotropous, usually (incompletely) bitegmic (sometimes unitegmic), incompletely tenuinucellar. Micropyle endostomal. Outer integument four to ten cell layers thick (in Hydrocera six to eight cell layers thick). Inner integument three to six cell layers thick. Micropylar epistase present. Megagametophyte usually monosporous, Polygonum type (sometimes disporous, 8-nucleate, Allium type), finally strongly prolonged. Endosperm development helobial (cellular?). Endosperm haustorium chalazal (in Hydrocera also micropylar). Embryogenesis onagrad.

Fruit Usually a loculicidal and septifragal carnose explosion capsule, dehiscing by capsule walls rolling in from base due to tensions in fleshy convex fruit wall; tensions liberated at slightest touch by bursting of wall along septa (in Hydrocera a drupaceous schizocarp with hard endocarp splitting into five mericarps each with one seed and two air chambers).

Seeds Seed pachychalazal. Seed mucilaginous hairs sclerotized, with spiral thickenings. Aril? Calciumoxalate raphide bundles abundant. Exotesta lignified, thickened. Mesotesta crushed. Endotesta? Tegmen? Testa in Hydrocera sclerotized, consisting of six to eight layers of thickened cells and five layers of unthickened cells. Perisperm not developed. Endosperm thin, rich in lipids, poor in starch. Embryo large, straight, well differentiated, without chlorophyll. Cotyledons two, large, planoconvex. Germination phanerocotylar.

Cytology n = (3–)7–10 (or more, up to 33; Impatiens), n = 8 (Hydrocera)

DNA Intron present in mitochondrial gene coxII.i3. Duplication of class B DEF genes.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin?), cyanidin, non-hydrolyzable tannins, proanthocyanidins (prodelphinidins), and naphthoquinones and their derivatives (e.g. lawsone) present. Ellagic acid, alkaloids, saponins, and cyanogenic compounds not found. Seed oils with glycerides of acetic acid and parinaric acid. Xyloclucans sometimes present in endosperm.

Distribution Madeira, East Asia to the Korean Peninsula, Japan and Taiwan, Hainan, Southeast Asia, Malesia to New Guinea, eastern and southeastern United States, Mexico, tropical America.

Fossils Two species of Clethra are known from the Miocene of Europe. In addition several fossilized capsules from Maastrichtian sediments in Europe have been assigned to Clethraceae, although their systematic affiliation may be questioned, such as Disoclethra with three species, Purdiaeopsis campanulatus and Valvaecarpus with four species. A fossil flower, Glandulocalyx upatoiensis, from the Santonian of Georgia has been attributed to Actinidiaceae or Clethraceae by Schönenberger & al. (2012).

Habit Bisexual (rarely functionally gynodioecious), evergreen or deciduous trees or shrubs.

Vegetative anatomy Arbuscular mycorrhiza present. Phellogen ab initio pericyclic. Stem in Clethra with prominent endodermis and heterogeneous medulla. Vessel elements with scalariform perforation plates (often with numerous cross-bars); lateral pits opposite or alternate, bordered pits. Vestured pits sometimes present. Imperforate tracheary xylem elements tracheids or fibre tracheids with bordered pits, non-septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty. Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace. Raphide cells absent.

Leaves Alternate (spiral), simple, entire, sometimes coriaceous, with conduplicate-subplicate ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate or annular; petiole with medullary bundle. Venation pinnate, eucamptodromous, brochidodromous or semicraspedodromous (in Purdiaea acrodromous to palinactinodromous). Stomata anomocytic, anisocytic, paracytic or actinocytic. Cuticular wax crystalloids? Domatia as hair tufts present in some species. Petiole and phloem with secretory cells. Calciumoxalate druses abundant. Leaf margins usually serrate or glandular serrate (sometimes entire).

Inflorescence Terminal or axillary, simple racemes, or paniculate, fasciculate or umbel-like cymose. Bracts often caducous (in Purdiaea large). Floral prophylls (bracteoles) absent.

Flowers Actinomorphic or slightly zygomorphic, small. Pedicel articulated. Hypogyny. Sepals five (or six), with imbricate quincuncial aestivation, persistent, free or partially connate, sometimes unequal in length. Petals five (or six), with imbricate quincuncial aestivation, caducous, usually free (rarely connate below). Ovary base often nectariferous. Disc absent.

Androecium Stamens usually 5+5 (rarely 6+6; in Purdiaea with outer stamens alternipetalous), diplostemonous (often secondarily obdiplostemonous). Filaments free from each other, usually free from tepals (sometimes adnate to petal bases). Anthers bilobate-sagittate to caudate (seemingly apically prolonged), ventrifixed (dorsifixed?), versatile, late inverted (finally resupinated), tetrasporangiate, at anthesis inverted from extrorse to introrse, poricidal (dehiscing by seemingly apical pores or short slits). Tapetum secretory, with binucleate cells. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolporate, shed as monads, bicellular at dispersal. Exine tectate, with granular infratectum, rugulate or psilate.

Gynoecium Pistil composed of three (to five) connate carpels. Ovary superior, usually trilocular (rarely quadri- or quinquelocular). Style single, simple. Stigma simple or usually trilobate (rarely quadri- or quinquelobate), papillate, Dry type. Pistillodium absent.

Ovules Placentation usually axile (in Purdiaea apical). Ovules few to numerous (in Purdiaea one) per carpel, usually anatropous (in Purdiaea orthotropous, pendulous), unitegmic, tenuinucellar. Integument ? cell layers thick (in Purdiaea gradually degenerating). Megagametophyte monosporous, Polygonum type. Endosperm development cellular. Endosperm haustoria chalazal and micropylar. Embryogenesis asterad.

Fruit Usually a loculicidal capsule, sometimes with persistent calyx (in Purdiaea a nut).

Seeds Aril absent. Seed coat often winged. Testa consisting of thin exotesta, with theoid exotestal thickenings (testa in Purdiaea indistinct, disappearing). Perisperm not developed. Endosperm copious, fleshy, oily and with hemicellulose. Embryo short, straight, well differentiated, chlorophyll? Cotyledons two, short. Germination phanerocotylar.

Phytochemistry Flavonols (quercetin), cyanidin, triterpenes, ellagic and gallic acids, and proanthocyanidins (prodelphinidin) present. Iridoids? Alkaloids and cyanogenic compounds not found. Carbohydrates stored as oligosaccharides with kestose or isokestose bonds.

Distribution Coastal plains in southeastern United States, the West Indies, Central America, northern South America.

Fossils Fossil wood and pollen grains assigned to Cyrilla are found in Oligocene layers in North America. Cenozoic brown coal in western Europe contains fossil wood similar to that in Cyrilla. The fossil fruits of Epacridicarpum from the Late Cretaceous and the Cenozoic of Europe have also been assigned to Cyrillaceae, but this systematic placement is doubtful.

Vegetative anatomy Phellogen ab initio pericyclic. Vessel elements with scalariform perforation plates; lateral pits scalariform or opposite, bordered pits. Imperforate tracheary xylem elements tracheids or fibre tracheids with bordered pits, non-septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse. Sieve tube plastids Pcf type (with protein crystals and fibrils, but no starch). Nodes 1:1, unilacunar with one leaf trace. Raphid cells absent. Wood ray cells sometimes with prismatic calciumoxalate crystals.

Leaves Alternate (spiral), simple, entire, sometimes coriaceous, with supervolute ptyxis. Stipules and leaf sheath absent; lateral or sublateral red or blackish ligulate? glandular structures – colleters – at buds and bracts may be reduced stipules. Petiole vascular bundle transection annular, complex or deeply concave. Venation pinnate. Stomata anomocytic. Cuticular wax crystalloids? Epidermis with or without mucilage cells. Mesophyll with calciumoxalate as druses and solitary prismatic crystals. Leaf margin entire.

Inflorescence Terminal or axillary, racemes. Floral prophylls (bracteoles) absent?

Flowers Actinomorphic (occasionally slightly zygomorphic). Hypogyny. Sepals five (to seven), with open (or imbricate quincuncial) aestivation, unequal in size, persistent, connate at base. Petals five (to seven), with imbricate quincuncial aestivation, free or connate at base, in Cyrilla nectariferous. Disc intrastaminal (in Cliftonia with nectaries?).

Androecium Stamens five (to seven), haplostemonous, antesepalous (inner whorl absent; Cyrilla), or (5+5) 6+6 or 7+7, diplostemonous (Cliftonia). Filaments free from each other and from tepals. Anthers dorsifixed, not inverted at anthesis, versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, with binucleate cells. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains 3(–6)-colporate, shed as monads, bicellular at dispersal. Exine tectate, with granular infratectum, microperforate, smooth.

Gynoecium Pistil composed of (two to) five connate carpels. Ovary superior, (bilocular to) quinquelocular. Basal part of ovary wall nectariferous? Style single, simple, very short, or absent. Stigma punctate or (bilobate to) quinquelobate, non-papillate, Dry type. Pistillodium absent.

Ovules Placentation axile or apical. Ovules one to three per carpel, anatropous, pendulous, usually apotropous, unitegmic, tenuinucellar. Integument four to seven cell layers thick, sometimes vascularized, gradually degenerating. Megagametophyte monosporous, Polygonum type (in at least Cliftonia sometimes also aposporous Hieracium type). Antipodal cells persistent. Endosperm development cellular. Endosperm haustoria chalazal and micropylar. Embryogenesis?

Fruit A one- to four-seeded dry drupe or a one-seeded two- to five-winged samara, often with persistent and accrescent calyx. Pericarp with tannins and druses.

Seeds Aril absent. Testa indistinct, early disappearing, with theoid exotestal thickenings? Perisperm not developed. Endosperm moderately developed, fleshy. Embryo large, straight, well differentiated, chlorophyll? Cotyledons two, small. Germination phanerocotylar?

Phytochemistry Insufficiently known. Flavonols (kaempferol, quercetin, myricetin), cyanidin, and ellagic acid (Cyrilla) present. Alkaloids and cyanogenic compounds not found. Condensed and hydrolyzable tannins? Iridoids?

Galacinaceae D. Don in Edinburgh New Philos. J. 6: 53. Oct-Dec 1828 [’Galacinae’]; Diapensiineae Link, Handbuch 1: 595. Jan-Aug 1829 [‘Diapensiaceae’]; Diapensiales Engl. et Gilg in Engler, Syllabus, ed. 9-10: 314. Nov-Dec 1924; Diapensianae Doweld, Tent. Syst. Plant. Vasc.: xlv. 23 Dec 2001

Distribution Cold-temperate and arctic-alpine regions on the Northern Hemisphere, the Himalayas, eastern Tibet, western China, northern Burma, Japan, Taiwan, southeastern United States.

Fossils Uncertain. Actinocalyx bohrii was described from the Late Cretaceous of southern Sweden. It had separate stylodia and the pollen grains were smaller than in extant Diapensiaceae. However, the systematic affiliation is questioned (Martínez-Millán 2010).

Vegetative anatomy Mycorrhiza ectotrophic and endotrophic. Phellogen ab initio usually pericyclic (sometimes superficial). Pericyclic fibres usually absent (present in Shortia). Vessel elements usually with simple (sometimes scalariform) perforation plates; lateral pits? Imperforate tracheary xylem elements ? with bordered pits. Wood rays absent. Axial parenchyma? Sieve tube plastids S type. Nodes usually 3:3, trilacunar with three leaf traces (in Pyxidanthera 1:1, unilacunar with one trace). Calciumoxalate crystals (single or compound) sometimes present.

Leaves Alternate (spiral), simple, entire, coriaceous, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate to annular; medullary bundles sometimes present. Venation pinnate to palmate or leaves uninerved, in Diapensia and Pyxidanthera inverted eucamptodromous (lateral veins curved at base, narrowing without fusing with main vein or other lateral veins), in Galax and Shortia actinodromous or camptodromous, in Berneuxia brochidodromous; secondary veins subpinnate to palmate. Stomata usually anomocytic, without subsidiary cells (rarely anisocytic). Cuticular wax crystalloids? Calciumoxalate as groups of crystals or single crystals (Galax). Leaf margin serrate or entire.

Inflorescence Terminal, raceme-like cymose, or flowers solitary terminal (sometimes axillary).

Flowers Actinomorphic. Hypogyny. Sepals five, with imbricate aestivation, persistent, free or connate into tube. Petals five, usually with imbricate (rarely contorted) aestivation, caducous, usually connate below (in Galax free or almost free), sometimes dentate at apex. Basal part of ovary in some species nectar-producing. Disc absent or strongly reduced.

Androecium Stamens five, haplostemonous, antesepalous, alternipetalous. Filaments flattened, usually free (in Galax connate at base), at least outer staminal whorl epipetalous. Anthers usually basifixed (or transverse with horizontal thecae), incurved (inverted during development, base becoming directed upwards), non-versatile, in Galax and Pyxidanthera with basal appendages, usually tetrasporangiate (in Galax disporangiate), introrse, longicidal (dehiscing by longitudinal [Berneuxia, Diapensia, Shortia] or transverse [Galax, Pyxidanthera] slits). Tapetum secretory, with binucleate cells. Staminodia absent in Diapensia and Pyxidanthera; in Berneuxia, Galax and Shortia five epipetalous antepetalous inner staminodia alternating with five fertile outer (alternipetalous) stamens.

Pollen grains Microsporogenesis simultaneous. Pollen grains tri- or hexacolp(oroid)ate, shed as monads, tricellular (bicellular?) at dispersal. Exine semitectate, with columellate infratectum, reticulate, or intectate.

Gynoecium Pistil composed of three connate carpels; median carpel adaxial. Ovary superior, trilocular, sometimes nectariferous. Style single, usually simple (in Diapensia trilobate), hollow (with stylar canal). Stigma usually single, shortly trifid (stigmas in Diapensia capitate), non-papillate, Wet type. Pistillodium absent.

Ovules Placentation usually axile (sometimes parietal). Ovules several to numerous per carpel, usually anatropous (sometimes hemianatropous or campylotropous), unitegmic, tenuinucellar. Integument five to seven cell layers thick. Megagametophyte monosporous, Polygonum type (in at least Pyxidanthera with large amount of starch). Synergids in Diapensia with a filiform apparatus. Antipodal cells sometimes proliferating (in Shortia up to c. 40 cells), persistent. Endosperm development cellular. Endosperm haustoria absent. Embryogenesis solanad?

Seeds Aril absent? Exotesta two or three cell layers thick, with thick inner cell walls, with theoid exotestal thickenings. Endotesta? Perisperm not developed. Endosperm copious, fleshy, containing lipids. Embryo relatively large, straight to somewhat curved, well differentiated, chlorophyll? Cotyledons two. Germination?

Phytochemistry Flavonols (kaempferol, quercetin, gossypetin), ellagic and gallic acids present. Ursolic acid, proanthocyanidins, alkaloids, and cyanogenic compounds not found. Iridoids? Aluminium accumulated.

Gürcke in Engler et Prantl, Nat. Pflanzenfam., IV, 1: 153. Dec 1891, nom. cons.

Guaiacanaceae Juss., Gen. Plant.: 155. 4 Aug 1789 [’Guiacanae’]; Diospyraceae Vest, Anleit. Stud. Bot.: 271, 294. 1818 [’Diospyroideae’]; Diospyropsida Brongn., Enum. Plant. Mus. Paris: xxi, 72. 12 Aug 1843 [’Diospyroideae’]; Diospyrales Prantl, Lehrb. Bot.: 216. 20 Apr 1874 [’Diospyrinae’]; Ebenales Engl., Syllabus: 155. Apr 1892; Ebenineae Bessey in C. K. Adams, Johnson’s Universal Cyclop. 8: 463. 15 Nov 1895 [‘Ebenales’]; Diospyrineae Engl., Syllabus, ed. 2: 170. Mai 1898; Lissocarpaceae Gilg in Engler et Gilg, Syllabus, ed. 9-10: 324. 6 Nov 1924, nom. cons.

Distribution Temperate to tropical regions on both hemispheres, with their highest diversity in Madagascar and Malesia; some species in temperate parts of eastern North America, India, eastern China, and Japan.

Fossils Leaves, flowers and fruits attributed to Ebenaceae are known from the Eocene onwards, also from Europe. Austrodiospyros cryptostoma is represented by male flowers and leaves from the Eocene of southeastern Australia. Diospyros pollen have been found in Eocene strata of North America.

Habit Usually dioecious (rarely monoecious, polygamomonoecious or bisexual), usually evergreen (rarely deciduous) trees or shrubs. Bark, roots and heartwood often black or, in air, blackening.

Vegetative anatomy Phellogen ab initio usually superficial (sometimes pericyclic). Primary medullary rays narrow. Vessel elements usually with simple (rarely scalariform) perforation plates; lateral pits alternate or opposite, bordered pits. Imperforate tracheary xylem elements fibre tracheids or libriform fibres with simple pits, non-septate. Wood rays uniseriate or multiseriate. heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty vasicentric, reticulate, or banded. Wood elements often storied. Sieve tube plastids S type. Nodes usually 1:1, unilacunar with one leaf trace (sometimes 1:3 or 3:3, unilacunar or trilacunar with three traces). Heartwood with gum-like substances. Sclereids present. Prismatic calciumoxalate crystals frequent.

Trichomes Hairs unicellular, simple or furcate, often malpighiaceous, with branches of unequal lengths, or multicellular, uniseriate or stellate (in Lissocarpa absent); glandular hairs, sometimes peltate-lepidote.

Leaves Usually alternate (spiral or distichous; rarely opposite or verticillate), simple, entire, coriaceous, with conduplicate ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate. Abaxial side of lamina usually with flattened glands, extrafloral nectaries. Venation pinnate. Stomata usually anomocytic (sometimes cyclocytic or actinocytic). Cuticular wax crystalloids absent (in Lissocarpa?). Mesophyll often with sclerenchymatous idioblasts containing brachysclereids, osteosclereids or other types of sclereids. Extrafloral nectaries often present on abaxial side. Secretory cavities absent. Leaf margin usually entire.

Inflorescence Axillary fasciculate, raceme-like or paniculate cymose (flowers sometimes solitary, terminal).

Flowers Actinomorphic. Pedicel articulated. Usually hypogyny (in Lissocarpa epigyny). Sepals three to five (to eight), with valvate or imbricate aestivation, persistent, connate. Petals three to five (to eight), with contorted aestivation, connate (corolla in Lissocarpa usually with eight-lobed corona). Nectariferous disc usually absent.

Androecium Stamens (three to) twelve to 20 (to c. 100), as many as or twice or four (or more) times the number of petals, usually unequal in length, in two (to four) whorls. Filaments flattened, free or connate two, three or more together (sometimes all filaments connate into a tube), usually adnate at base to corolla tube. Anthers basifixed, non-versatile, tetrasporangiate, external anthers introrse, mid-anthers latrorse and internal anthers extrorse, usually longicidal (dehiscing by longitudinal slits; sometimes poricidal, with apical pores); connective often slightly prolonged. Tapetum secretory, with multinucleate cells. Female flowers usually with a single staminodial whorl.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolporate (rarely tetracolporate; in Lissocarpa triporate), usually shed as monads (rarely tetrads), bicellular at dispersal. Exine tectate, with granular infratectum (Diospyros), rugulate, microrugulate, striate, gemmate or granulate (in Lissocarpa fossulate to rugulate).

Gynoecium Pistil composed of two to eight connate carpels. Ovary usually superior (in Lissocarpa inferior), usually bilocular to 16-locular; each locule usually divided by secondary septa (ovary rarely unilocular; ovary in Lissocarpa quadrilocular, not divided by secondary septa). Stylodia two to eight, free or partially connate (style in Lissocarpa single, simple, clavate). Stigmas capitate or bilobate (stigma in Lissocarpa often slightly quadrilobate), non-papillate, Dry type. Male flowers usually with pistillodium.

Ovules Placentation usually apical (in Lissocarpa axile). Ovules one or two per carpel, anatropous, pendulous, apotropous, bitegmic, tenuinucellar. Micropyle bistomal or endostomal. Outer integument three to seven cell layers thick. Inner integument five to ten cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development cellular (nuclear?). Endosperm haustoria? Embryogenesis probably chenopodiad.

Fruit Usually a berry (rarely capsule-like) with persistent and usually accrescent calyx. Pericarp with tanniniferous idioblasts.

Seeds Aril absent. Testa vascularized, with circumperipheral vascular loop surrounding seed. Tegmen? Perisperm not developed. Endosperm copious, hard, oily, with mannose-rich polysaccharides, sometimes ruminate. Embryo straight or curved, well differentiated (Lissocarpa), without chlorophyll. Cotyledons two, foliaceous. Radicula long. Germination phanerocotylar or cryptocotylar.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), cyanidin, C-30-oxidized triterpenes, oleanolic acid derivatives, ellagic and gallic acids, methylated ellagic acids, condensed tannins, proanthocyanidins (prodelphinidins), saponins, naphthoquinone derivatives of 7-methyljugone and plumbagin (droserone; bark and wood blackening when substances oxidized, cf. ebony), anthraquinones, and benzopyrones present. Alkaloids and cyanogenic compounds rare. Aluminium accumulated in some species.

Kalmiaceae Durande, Notions Elém. Bot.: 271. 1782 [’Kalmiae’], nom. illeg.; Rhododendraceae Juss., Gen. Plant.: 158. 4 Aug 1789 [’Rhododendra’]; Rhodoraceae Vent., Tabl. Règne Vég. 2: 449. 5 Mai 1799; Ledaceae J. F. Gmel., Allg. Gesch. Pflanzengifte, ed. 2: 404. Jun 1803 [’Leda’]; Epacridaceae R. Br., Prodr. Fl. Nov.-Holl.: 535. 27 Mar 1810 [’Epacrideae’], nom. cons.; Azaleaceae Vest, Anleit. Stud. Bot.: 272, 294. 1818 [’Azaleoideae’]; Monotropaceae Nutt., Gen. N. Amer. Pl. 1: 272. 14 Jul 1818 [’Monotropeae’], nom. cons.; Vacciniaceae DC. ex Perleb, Vers. Artzneikr. Pfl.: 228. Mai 1818 [’Vaccinieae’], nom.cons.; Empetrales Bercht. et J. Presl, Přir. Rostlin: 251. Jan-Apr 1820 [‘Empetreae’]; Epacridales R. Br. ex Bercht. et J. Presl, Přir. Rostlin: 251. Jan-Apr 1820 [‘Epacrideae’]; Monotropales Bercht. et J. Presl, Přir. Rostlin: 252. Jan-Apr 1820 [‘Monotropeae’]; Empetraceae Hook. et Lindl. in W. J. Hooker, Fl. Scot.: 297. 10 Mai 1821 [’Empetreae’], nom. cons.; Pyrolaceae Lindl., Syn. Brit. Fl.: 175. 16 Mar 1829 [’Pyroleae’], nom. cons.; Empetrineae Link, Handbuch 1: 617. 4-11 Jul 1829 [‘Empetreae‘]; Epacridineae Link, Handbuch 1: 601. 4-11 Jul 1829 [‘Epacrideae‘]; Ericineae Link, Handbuch 1: 601. 4-11 Jul 1829 [‘Ericeae genuinae‘]; Vacciniales Dumort., Anal. Fam. Plant.: 28. 1829 [‘Vaccinarieae’]; Stypheliaceae Horan., Prim. Lin. Syst. Nat.: 72. 2 Nov 1834 [’Stypheliaceae (Epacrideae)’]; Arbutaceae (Meisn.) Miers in Bromhead in Mag. Nat. Hist., n.s., 4: 337, 338. Jul 1840 [’Arbuteae’]; Andromedaceae Döll, Rhein. Fl.: 428. 24-27 Mai 1843 [’Andromedeae’]; Arbutineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.]: 987. 1846; Pyrolineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.]: 987, 989. 1846 [‘Pyroleae‘]; Rhododendrineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.]: 987, 996. 1846 [‘Rhododendreae‘]; Rhodorales Horan., Char. Ess. Fam.: 105. 30 Jun 1847 [‘Rhodorastra’]; Hypopityaceae Klotzsch in Linnaea 24: 11. Mai 1851 [’Hypopithieae’]; Menziesiaceae Klotzsch in Linnaea 24: 11. Mai 1851; Siphonandraceae Klotzsch in Linnaea 24: 11. Mai 1851, nom. illeg.; Arctostaphylaceae J. Agardh, Theoria Syst. Plant.: 106. Apr-Sep 1858 [’Arctostaphyleae’]; Salaxidaceae J. Agardh, Theoria Syst. Plant.: 104. Apr-Sep 1858 [’Salaxideae’]; Diplarchaceae Klotzsch in Monatsber. Königl. Preuss. Akad. Wiss. Berlin 1859: 15. 1857 [’Diplarchaceen’], nom. illeg.; Oxycoccaceae A. Kern., Pflanzenleben 2: 713, 714. 8 Aug 1891; Prionotaceae Hutch., Evol. Phylog. Fl. Pl.: 306. 28 Aug 1969

Distribution Cosmopolitan, although few species in tropical lowland regions, with their largest diversity in the Himalayas to southwestern China, New Guinea, South Africa, Australia, and New Zealand.

Fossils Seeds and fruits from several genera with only extra-European recent distribution in the Northern Hemisphere, e.g. Eubotrys, Leucothoe, Lyonia and Zenobia, have been found in Late Cretaceous and Early Palaeogene strata in Europe. Diplycosiopsis walbeckensis and Viticocarpum minimum from the Maastrichtian of Germany is represented by drupes with five many-seeded and one-seeded locules, respectively. It is, however, uncertain whether they emanate from species of Ericaceae. Fossil leaves and pollen grains from supposed Styphelioideae are known from (possibly the Late Cretaceous to) the Early Palaeogene onwards in, above all, Australia.

Habit Usually bisexual (rarely monoecious or dioecious), usually evergreen (sometimes deciduous) shrubs or suffrutices (rarely trees, lianas or perennial herbs). Some genera consist of achlorophyllous and mycotrophic plants. Numerous species are xerophytic. Some species are helophytes. A starchy lignotuber is present especially in many species in mediterranean climates.

Vegetative anatomy Ectomycorrhiza usually with Basidiomycotina (i.a. Sebacinales) or often with certain Ascomycotina (particularly Pezizales, e.g. Hymenoscyphus) as intracellular hyphal complexes in root epidermis (monotropoid, ericoid or arbutoid mycorrhiza); ericoid mycorrhiza penetrates only epidermal and outer cortical cells of hair roots (consisting of endodermis, exodermis, tracheids, sieve tubes and companion cell) covered with hyphae (Hartig net and fungal mantle not formed); arbutoid mycorrhiza penetrates epidermal and outer cortical cells of hair roots, these being covered by Hartig net with hyphae following cell walls or forming complete envelope (Hartig net and fungal mantle formed); in monotropoid mycorrhiza root covered in the same way, although hyphae form peg-shaped invaginations into exodermal hair root cells (Hartig net and fungal mantle formed); arbuscular mycorrhiza present in Enkianthus; standard ectomycorrhiza not known in Ericaceae. Endophytic fungi frequently present. Phellogen ab initio usually pericyclic (sometimes superficial; absent in Monotropoideae). Pericyclic fibres little developed. Secondary lateral growth usually normal (absent in most Monotropoideae). Vessel elements usually with simple or scalariform (rarely reticulate) perforation plates; lateral pits scalariform, opposite or alternate (vessels absent in most Monotropoideae), simple and/or bordered pits. Vestured pits sometimes present. Imperforate tracheary xylem elements tracheids or fibre tracheids or libriform fibres with simple and/or bordered pits, usually non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, usually heterocellular. Axial parenchyma apotracheal diffuse (rarely diffuse-in-aggregates), or paratracheal scanty vasicentric or unilateral, or absent. Sieve tube plastids usually S type (sometimes S₀ type). Endodermis rarely with suberised cell walls. Nodes usually 1:1, unilacunar with one leaf trace (rarely ≥3:≥3, trilacunar or multilacunar with three or more traces). Cortex with sclereids. Raphid cells absent. Calciumoxalate druses or single prismatic crystals often frequent.

Trichomes Hairs unicellular or multicellular, uniseriate or branched, dendritic, stellate, lepidote, bristle-like, sometimes candelabra-, funnel- or cup-shaped; glandular hairs present (sometimes peltate-lepidote), often thick and colleter-like.

Leaves Alternate (sometimes distichous or tetrastichous), opposite or verticillate, simple, entire, usually coriaceous, often ericoid (in Monotropoideae usually scale-like), with convolute, involute or revolute ptyxis. Stipules and leaf sheath absent. Colleter-like thick glandular hairs often present on adaxial side of petiole base. Petiole vascular bundle transection usually arcuate (sometimes annular). Extrafloral nectaries sometimes present on petiole or base of lamina. Venation usually pinnate (rarely palmate or parallelodromous) or leaves one-veined. Stomata usually anomocytic or paracytic (rarely parallelocytic, cyclocytic or stephanocytic). Cuticular wax crystalloids usually amorphous or as platelets (sometimes as tubuli, dominated by β-diketones, or as coiled or triangular rodlets). Epidermis with or without mucilage cells. Secretory oil cavities present or absent. Leaf margin serrate, crenate or entire; teeth united via multicellular hairs.

Inflorescence Terminal or axillary, racemes, spikes, panicles or fascicles (sometimes umbels, rarely heads or flowers solitary).

Flowers Actinomorphic or zygomorphic. Hypogyny to epigyny. Sepals (two to) four or five (to seven), with imbricate quincuncial aestivation, persistent, usually connate at base. Petals (three or) four or five (to seven), with valvate or imbricate (rarely contorted) aestivation, persistent or caducous, usually connate into campanulate, tubular or urceolate corolla (rarely free; in Richea connate into calyptra). Nectary or nectariferous disc (annular or of separate parts, intrastaminal) usually at base (sometimes at apex) of ovary (sometimes absent).

Androecium Stamens usually four or five (rarely two or three), antesepalous, alternipetalous, or 4+4, 5+5(–16), usually obdiplostemonous. Filament usually free from each other and from tepals (in, e.g., Styphelioideae often epipetalous). Anthers basifixed or dorsifixed, late inverted, finally resupinate (in Enkianthoideae, Monotropoideae and Arbutoideae sometimes not inverted), often versatile, usually free (rarely connate), often with two (rarely four) small usually apical horn-shaped outgrowths (sometimes dorsal spurs; in Vaccinioideae sometimes tubular outgrowths), usually tetrasporangiate (in Styphelioideae disporangiate), subintrorse and poricidal (dehiscing by apical pores), or introrse (rarely latrorse or extrorse) and longicidal (dehiscing by longitudinal slits; in Styphelioideae usually with a single slit). Tapetum secretory, with uni-, bi- or multinucleate cells. Staminodia absent. Secondary pollen display present in Styphelioideae.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually 3(–5)-colporate (sometimes 3(–5)-colpate, 3(–5)-colporoidate or porate), usually shed as tetrahedral tetrads (sometimes as monads, rarely as dyads, triads or polyads; in some Styphelioideae three pollen cells of tetrad degenerate), usually bicellular (sometimes tricellular) at dispersal. Exine tectate, with granular infratectum, scabrate, verrucate, rugulate, microrugulate or psilate. Viscinous threads present on pollen surface in many groups.

Gynoecium Pistil composed of (one to) four or five (to twelve) connate, usually antepetalous (in, e.g., Monotropa, Dracophyllum and Vaccinium antesepalous) carpels. Ovary superior to inferior, (unilocular to) quadri- or quinquelocular, nectariferous at base. Style single, simple, usually hollow (with stylar canal; rarely widened at apex). Stigma capitate or punctate to somewhat lobate or peltate, usually papillate, Dry or Wet type. Male flowers rarely with pistillodium.

Ovules Placentation usually axile to intrusively parietal (parietal placentation present in, e.g., Monotropoideae; rarely apical or basal). Ovules usually numerous (sometimes several, rarely single) per carpel, anatropous or hemianatropous to (almost) campylotropous, pendulous to ascending, apotropous to epitropous, unitegmic, tenuinucellar. Integument two to five cell layers thick. Megagametophyte usually monosporous, Polygonum type (in Arbutoideae sometimes disporous, Allium type). Synergids sometimes with a filiform apparatus. Antipodal cells often persistent, sometimes proliferating. Endosperm development cellular or nuclear. Endosperm haustoria chalazal and/or micropylar (rarely absent). Embryogenesis usually solanad (in Pyroleae caryophyllad).

Fruit A loculicidal or septicidal (rarely septifragal) capsule, a berry, or a drupe (with one or more pyrenes; rarely a nut surrounded by fleshy perianth), with persistent calyx.

Seeds Aril absent. Seed coat winged or unwinged. Testa usually one-layered, consisting of exotesta, with inner and radial but no outer cell walls thickened, with theoid exotestal thickenings (sometimes mucilaginous). Perisperm not developed. Endosperm copious, fleshy, often oily. Embryo straight, usually well differentiated (in Monotropoideae usually undifferentiated; in Monotropa uniflora two-celled, in M. hypopitys nine-celled, in Allotropa and Pleuricospora two- or three-celled, in Monotropsis four-celled, in Sarcodes several-celled with a globular structure, in Chimaphila undifferentiated), with or without chlorophyll. Cotyledons two, short. Germination phanerocotylar.

Cytology n = 11 (Enkianthoideae); n = 8, 13, 19, 23 etc. (Monotropoideae); n = 13 (Arbutoideae); n = 13 (Cassiopoideae); n = 12, 13 (26) (Ericoideae); n = 16 (Harrimanelloideae); n = 12 (17, 18, 24) (Vaccinioideae); n = 4, 6–13 (Styphelioideae) – Polyploidy occurring (12x in Rhododendron manipurense). Protein crystals present in nucleus in some species.

DNA Plastid gene infA lost/defunct (Andromeda, Chamaedaphne, Pyrola). Mitochondrial intron coxII.i3 lost. Mitochondrial coxI intron present in Pyrola.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin, gossypetin), cyanidin, catechins, Route I secoiridoids (also decarboxylated iridoids?), Group I carbocyclic iridoids (aucubin, daphylloside, monotropein), Group IV carbocyclic iridoids (unedoside, a C8 iridoid glycoside, in Arbutus and Arctostaphylos), diterpenes (e.g. andromedotoxin in Arbutoideae and Ericoideae), triterpene acids (ursolic acid), ellagic acid (rare), gallic acid, ellagitannins (geraniin), proanthocyanidins (prodelphinidins), alkaloids, cyanogenic compounds, chlorogenic acid, 3-galactoside, benzo- and naphthoquinones, myo-inisitol, and ethereal oils present. Arbutin present in Arbutoideae, Monotropoideae and Vaccinioideae. Monotropeoside present in Monotropoideae. Saponins not found. Polysaccharides stored as oligosaccharides with kestose and isokestose bonds.

Use Ornamental plants, fruits (of Vaccinium, Oxycoccus, Myrtillus, Gaylussacia, Arctostaphylos, etc.), honey (from, e.g., Calluna vulgaris), timber, carpentries, smoking-pipes (from Erica arborea and E. scoparia), medicinal plants.

Systematics Ericaceae are sister to Cyrillaceae.

A probable topology is the following: [Enkianthoideae+[Monotropoideae+[Arbutoideae+[[Cassiopoideae+Ericoideae]+[Harrimanelloideae+[Styphelioideae+Vaccinioideae]]]]]] (Kron & al. 2002).

According to the ICBN, the name Ericaceae Juss. should be preferred before Vacciniaceae Adans.

Cladogram (simplified) of Ericaceae based on morphology and DNA sequence data (Kron & al. 2002). Monotropoideae are sister-group to Arbutoideae and Pyroloideae are sister to the remaining Ericaceae except Enkianthoideae, Monotropoideae and Arbutoideae (Rose & al. 2018).

Enkianthoideae (P. F. Stevens) Kron, Judd & Anderb. in K. A. Kron et al., Bot. Rev. (Lancaster) 68: 401. 23 Oct 2002

1/17. Enkianthus (15–17; Japan, southern China, northern Burma, Indochina). – Mycorrhiza arbuscular, endomycorrhizal, Paris type; hair roots absent. Medulla heterocellular, with small thick-walled lignified larger and thin-walled cells mixed. Vestured pits present. Hairs with distal end dying off long before proximal end. Leaves pseudoverticillate. Buds perulate (axillary branches developing exclusively in axils of perulae; also present in some Rhodoreae). Anthers with paired bristles and fibrous endothecium. Pollen grains shed as monads, tricellular at dispersal. Tectum with granulate surface. Ventral carpel bundles present in septal plane. Megagametophyte inserted on megasporangial pedestal. Megagametophyte with ear-like appendages. Seed raphe with vascular bundles. n = 11. – Enkianthus has the most well preserved remnants of pit membranes in Ericaceae.

[Monotropoideae+[Arbutoideae+[[Cassiopoideae+Ericoideae]+[Harrimanelloideae+[Styphelioideae+Vaccinioideae]]]]]

Mycorrhiza types of ectendomycorrhiza. Hair roots present, consisting of endodermis, exodermis, tracheid, sieve tube and companion cell. Fungal hyphae with complex coiled intrusions into hair root exodermal cells (only pegs present in Monotropeae and Pterosporeae). Anthers poricidal, with exothecium. Pollen grains shed in tetrahedral tetrads. Seed without chalazal vascular bundle. Insertions present in plastid gene matK.

Monotropoideae Arn., Botany: 118. 9 Mar 1832 [‘Monotropeae’]

14/c 50. Pyroleae Dumort., Anal. Fam. Plant.: 47. 1829. Pyrola (c 35; temperate regions on the Northern Hemisphere to mountains on Taiwan and northern Sumatra, temperate South America), Orthilia (1; O. secunda; temperate and arctic-alpine regions on the Northern Hemisphere), Chimaphila (5; C. japonica, C. maculata, C. menziesii, C. monticola, C. umbellata; Europe, temperate Asia, North and tropical America), Moneses (1; M. uniflora; temperate regions on the Northern Hemisphere). – Monotropeae Dumort., Anal. Fam. Plant.: 47. 1829. Allotropa (1; A. virgata; western United States), Cheilotheca (3; C. khasiana, C. malayana, C. sleumerana; Assam to West Malesia), Hemitomes (1; H. congestum; western United States), Monotropa (2; M. hypopitys, M. uniflora; temperate regions on the Northern Hemisphere, mountains in Central America), Monotropastrum (2; M. humile, M. sciaphilum; East Asia, Sumatra), Monotropsis (1; M. odorata; southeastern United States), Pityopus (1; P. californica; western United States), Pleuricospora (1; P. fimbriolata; southwestern Canada, western United States). – Pterosporeae Baill., Hist. Plant. 11: 161, 206. Jun-Jul 1891. Pterospora (1; P. andromeda; southern Canada, western and northeastern United States, northern Mexico), Sarcodes (1; S. sanguinea; western United States, Mexico). – Temperate and arctic-alpine regions on the Northern Hemisphere, mountains in Central America and mountains on Taiwan and northern Sumatra. Many species achlorophyllous and hyperparasitic mycoheterotrophs. Herbs with ericoid or arbutoid mycorrhiza (Pyroleae) or monotropoid mycorrhiza (with fungal hyphae forming peg-shaped invaginations into exodermal hair root cells; Monotropeae, Pterosporeae). Association between achlorophyllous Monotropoideae and mycorrhiza-forming basidiomycetes often specific. Hartig net frequently present. Sieve tube plastids at least in some achlorophyllous species S₀ type. Multicellular hairs usually absent (present in Pterosporeae). Leaves in Monotropeae and Pterosporeae scale-like, in Pyroleae foliaceous. Inflorescence racemose. Floral prophylls (bracteoles) usually absent (present in Pterospora and Sarcodes). Sepals and petals three to eight, often free (sometimes absent). Anthers with various types of dehiscence, usually longicidal (dehiscing by slits). Tapetum cells binucleate. Pollen grains usually shed as monads (in Pyroleae as tetrahedral tetrads). Placentation axile to intrusively parietal. Seed coat often winged. Testa with thickened or unthickened cell walls. Embryo reduced, small to minute and undifferentiated (Monotropa uniflora has bicellular embryo). n = 8, 11, 13, 16, 19, 23 (26). Cell nuclei with protein crystals. Polysaccharides stored as kestose and isokestose oligosaccharides. – Pyroleae (Pyroloideae Beilschm. in Flora 16(Beibl. 7): 72, 109. 14 Jun 1833) are sister-group to [Monotropeae+Pterosporeae]. Pyroleae are herbaceous, rhizomatous and have photosynthesizing foliaceous leaves. The mycorrhiza is ericoid or arbutoid. The anthers have very short or no tubules and no outgrowths. The pollen grains are usually shed as tetrahedral tetrads (sometimes as monads). Nectary is usually absent. The placentation is intrusively parietal. The testal walls are thin. A possible although unstable topology is [[Orthilia+Pyrola]+[Moneses+Chimaphila]] (Freudenstein & al. 1999; Liu & al. 2011). An alternative topology was recovered by Freudenstein & al. (2016), where Pyroleae (Pyroloideae) were sister-group to either [Arbutoideae+[Monotropeae+Pterosporeae]] or to core Ericaceae (with Arbutoideae sister to [Monotropeae+Pterosporeae]). The latter hypothesis was supported by Rose & al. (2018).

[Arbutoideae+[[Cassiopoideae+Ericoideae]+[Harrimanelloideae+[Styphelioideae+Vaccinioideae]]]]

Petiole vascular bundle transection annular. Floral prophylls (bracteoles) present. Anthers with poricidal dehiscence. Endothecium absent. Pollen grains shed as tetrahedral tetrads.

Arbutoideae (Benth.) Nied. in Engl. Bot. Jahrb. Syst. 11: 135. 18 Jun 1889

3–5/c 85. Arbutus (12; western Europe, the Canary Islands, the Mediterranean, southwestern Canada, western United States, northwestern Mexico; possibly paraphyletic and incl. Arctostaphylos, Comarostaphylis, Ornithostaphylos?), Arctostaphylos (c 60; southwestern Canada and western United States, Mexico, mountains in Central America; in Arbutus?), Arctous (3; A. alpina, A. microphyllus, A. ruber; temperate and arctic-alpine regions on the Northern Hemisphere), Comarostaphylis (10; southern California, northwestern Mexico; in Arbutus?), Ornithostaphylos (1; O. oppositifolia; southern California, northwestern Mexico; in Arbutus?). – Temperate regions on the Northern Hemisphere southwards to mountains in Central America, with their highest diversity in California. Mycorrhiza arbutoid. Hartig net frequently present. Corolla urceolate, with unicellular hairs on adaxial side. Filaments distinctly asymmetrically widened. Anthers with paired bristles. Endothecium absent. Style continuous. Ovules ten or fewer per carpel. Fruit a berry or a drupe (sometimes covered with large fleshy multicellular papillae), with bony or fibrous endocarp. Testal cell walls fairly thick. x = 13. C-8 iridoid glucosides and ellagic acid present.

[[Cassiopoideae+Ericoideae]+[Harrimanelloideae+[Styphelioideae+Vaccinioideae]]]

Mycorrhiza ericoid, usually with ascomycetes. Hartig net absent. Mycorrhiza-forming fungi usually ascomycetes. Stem with well developed pericyclic fibres connected to foliar vascular bundles. Anthers early inverted. Endothecium absent. Toxic andromedane diterpenes sometimes present.

[Cassiopoideae+Ericoideae]

Leaves opposite, ericoid (usually needle-like), with revolute ptyxis.

Cladogram of Cassiopoideae and Ericoideae based on DNA sequence data (Kron 1997).

Cassiopoideae (H. T. Cox ex P. F. Stevens) Kron et Judd in K. A. Kron et al., Bot. Rev. (Lancaster) 68: 404. 23 Oct 2002

1/12. Cassiope (12; arctic-alpine regions on the Northern Hemisphere, the Himalayas). – Mycorrhiza ericoid. Medulla Calluna type, with large thin-walled cells surrounded by smaller thick-walled lignified cells. Special fasciculate hairs present. Leaves decussate, linear, peltate, hypoascidiate etc., with flat to recurved ptyxis, without or with poorly developed associated fibres in mid vein. Inflorescence axillary, reduced. Bud scales absent. Anthers with bristles. Megagametophyte disporous.

Ericoideae Arn., Botany: 118. 9 Mar 1832 [’Ericeae’]

16/1.770–1.780. Phyllodoceae Drude in Engler et Prantl, Nat. Pflanzenfam. IV, 1: 31, 38. Oct 1889.Bejaria (15; southern United States, Mexico, Central America, tropical South America), Elliottia (4; E. bracteata, E. paniculata: Japan; E. pyroliflora: Alaska, western Canada, northwestern United States; E. racemosa: southeastern United States), Kalmia (10; Canada, United States, one species, K. ericoides, on Cuba, one species, K. procumbens, in arctic-alpine regions on the Northern Hemisphere), Epigaea (3; E. gaultherioides: eastern Turkey, the Caucasus; E. asiatica: Japan; E. repens: eastern United States), Rhodothamnus (2; R. chamaecistus: eastern Alps; R. sessilifolius: northeastern Turkey), Kalmiopsis (1; K. leachiana; Oregon), Phyllodoce (7; P. aleutica, P. breweri, P. caerulea, P. deflexa, P. empetriformis, P. glanduliflora, P. nipponica; cold-temperate and arctic-alpine regions on the Northern Hemisphere). – Bryantheae Gillespie et Kron in Brittonia 64(1): 75. 1 Mar 2012. Bryanthus (1; B. gmelinii; Kamchatka, Japan), Ledothamnus (7; L. atroadenus, L. decumbens, L. guyanensis, L. jauaensis, L. luteus, L. parviflorus, L. sessiliflorus; the Guayana Highlands). – Ericeae DC. ex Duby, Bot. Gall. 1: 316. 12-14 Apr 1828 [‘Ericaceae’]. Daboecia (2; D. cantabrica: Ireland to Spain; D. azorica: the Azores), Calluna (1; C. vulgaris; Europe, Turkey), Erica (c 860; Europe, Macaronesia, the Mediterranean, tropical and southern Africa, Madagascar, the Comoros, with their highest diversity in the Cape Provinces). – Empetreae Horan., Char. Ess. Fam.: 109. 17 Jun 1847. Ceratiola (1; C. ericoides; southeastern United States), Corema (2; C. album: Portugal, western Spain, the Azores; C. conradii: eastern Canada, northeastern United States), Empetrum (3; E. eamesii, E. nigrum, E. rubrum; temperate and arctic-alpine regions on the Northern Hemisphere, southern Andes in Chile from 35°S to 55°S, the Falkland Islands, Tristan da Cunha). – Rhodoreae DC. ex Duby, Bot. Gall. 1: 318. 12-14 Apr 1828 [‘Rhodoraceae’].Rhododendron (c 850; temperate and alpine regions on the Northern Hemisphere and south to mountains in Southeast Asia, Malesia to New Guinea and eastern Australia from northeastern Queensland to eastern Victoria, with their largest diversity in the Himalayas, western China, northern Burma and alpine New Guinea). – Temperate, arctic-alpine and montane regions in the Northern Hemisphere, mountain areas in the Southern Hemisphere, the Falkland Islands, Tristan da Cunha, with their highest diversity in South Africa, mountains in Southeast Asia and New Guinea. Usually bisexual (in Empetreae usually monoecious or dioecious). Mycorrhiza ericoid. Leaves alternate (spiral) or opposite (in Empetreae and Ericeae often verticillate), with usually flat (sometimes convolute, in Empetreae and Ericeae strongly revolute) ptyxis. Pedicel sometimes articulated. Petals usually four or five (in Rhodoreae often more than five; in Empetreae four, free). Anthers usually without appendages (in Ericeae often spurred). Pollen grains in Rhodoreae often with viscin threads. Stigma in Empetreae expanded. Ovule in Empetreae one per carpel, basal, apotropous. Fruit usually a septicidal capsule (in Erica also loculicidal; in Empetreae a drupe). Gossypetin present in Rhodoreae and Empetreae. – Phyllodoceae are sister to the remaining Ericoideae, according to Gillespie & Kron (2012), whereas Bryantheae are sister-group to the clade [Ericeae+[Empetreae+Rhodoreae]].

[Harrimanelloideae+[Styphelioideae+Vaccinioideae]]

Fruit with persistent and non-withering calyx.

Harrimanelloideae Kron et Judd in K. A. Kron et al., Bot. Rev. (Lancaster) 68: 409. 23 Oct 2002

1/2. Harrimanella (2; H. hypnoides: arctic-alpine regions in northern Europe to northern Ural, arctic Canada to Quebec and northeastern United States; H. stelleriana: high mountains in Japan, the Kurile Islands, Kamchatka, Alaska, northwestern Canada, northwestern United States). – Mycorrhiza ericoid? Leaves acicular, sublinear. Leaf margin entire. Flowers solitary, axillary. Floral prophylls (bracteoles) absent. Anthers spurred, poricidal.

[Styphelioideae+Vaccinioideae]

Cladogram (simplified) of Styphelioideae based on DNA sequence data (Crayn & Quinn 2000).

Styphelioideae Sweet, Fl. Australas.: ad t. 47. 1 Mai 1828 [‘Stypheliae’]

36/510–525. Prionoteae Drude in Engler et Prantl, Nat. Pflanzenfam. IV, 1: 72. Dec 1889. Lebetanthus (1; L. myrsinites; southernmost Andes, Patagonia, Tierra del Fuego), Prionotes (1; P. cerinthoides; Tasmania). – Archerieae Crayn et Quinn in Austral. Syst. Bot. 11: 23. 31 Mar 1998. Archeria (6; A. comberi, A. eriocarpa, A. hirtella, A. serpyllifolia: Tasmania; A. racemosa, A. traversii: New Zealand). – Oligarrheneae Crayn et Quinn in K. A. Kron et al., Bot. Rev. (Lancaster) 68: 412. 23 Oct 2002. Needhamiella (1; N. pumilio; southwestern Western Australia), Oligarrhena (1; O. micrantha; southwestern Western Australia), Dielsiodoxa (3; D. leucantha, D. oligarrhenoides, D. tamariscina; southwestern Western Australia). – Cosmelieae Crayn et Quinn in K. A. Kron et al., Bot. Rev. (Lancaster) 68: 413. 23 Oct 2002.Andersonia (c 35; southwestern Western Australia), Cosmelia (1; C. rubra; southwestern Western Australia), Sprengelia (4; S. distichophylla, S. incarnata, S. monticola, S. sprengelioides; eastern South Australia to southeastern Queensland, Tasmania). – Richeeae Crayn et Quinn in K. A. Kron et al., Bot. Rev. (Lancaster) 68: 412. 23 Oct 2002. Dracophyllum (50–55; eastern Queensland, eastern New South Wales, Tasmania, New Caledonia, New Zealand), Richea (11; southeastern New South Wales, Victoria, Tasmania, with their highest diversity in Tasmania), Sphenotoma (6; S. capitatum, S. dracophylloides, S. drummondii, S. gracilis, S. parviflorum, S. squarrosa; southwestern Western Australia). – Epacrideae Dumort., Anal. Fam. Plant.: 28. 1829 [‘Epacreae’]. Epacris (55–60; southeastern South Australia to southeastern Queensland, Tasmania, New Caledonia, New Zealand), Lysinema (6; L. ciliatum, L. conspicuum, L. elegans, L. fimbriatum, L. lasianthum, L. pentapetalum; southwestern Western Australia), Woollsia (1; W. pungens; southeastern Queensland, eastern New South Wales). – Styphelieae Bartl., Ord. Nat. Plant.: 158. Sep 1830 [‘Stypheliae’]. Pentachondra (4; P. dehiscens, P. ericifolia, P. involucrata, P. pumila; eastern New South Wales, eastern Victoria, Tasmania, New Zealand), Acrothamnus (6; A. colensoi, A. hookeri, A. maccraei, A. montanus, A. spathaceus, A. suaveolens; northeastern Queensland, eastern New South Wales, eastern Victoria, Tasmania, New Zealand), Acrotriche (16; southwestern Western Australia, southeastern South Australia to eastern Queensland, Tasmania), Androstoma (2; A. verticillata: Tasmania; A. empetrifolia: New Zealand), ’Astroloma’ (28; southwestern Western Australia, southeastern South Australia, Victoria, eastern New South Wales, Tasmania; non-monophyletic), Stenanthera (5; S. brachyloma, S. ciliata, S. conostephioides, S. pinifolia, S. squamuligera; eastern New South Wales, Victoria, Tasmania), Brachyloma (8; southwestern Western Australia, southeastern South Australia to southeastern Queensland, Tasmania), Coleanthera (3; C. coelophylla, C. myrtoides, C. virgata; southwestern Western Australia), Conostephium (10–12; southwestern Western Australia), Croninia (1; C. kingiana; southwestern Western Australia), Cyathodes (5; C. dealbata, C. glauca, C. petiolaris, C. platystoma, C. straminea; Tasmania), Cyathopsis (1; C. floribunda; New Caledonia), Decatoca (1; D. spenceri; New Guinea), Leptecophylla (13; New Guinea, Tasmania, New Zealand to the Hawaiian Islands), ’Leucopogon’ (155–165; Malesia to New Guinea, Australia, Tasmania, New Caledonia, New Zealand; non-monophyletic), ’Lissanthe’ (7; L. brevistyla, L. pluriloculata, L. powelliae, L. rubicunda, L. sapida, L. scabra, L. strigosa; southwestern Western Australia, southeastern South Australia to southeastern Queensland, Tasmania; non-monophyletic), Melichrus (13?; M. adpressus, M. erubescens, M. hirsutus, M. procumbens, M. urceolatus; continental southwestern Western Australia, eastern Queensland to Victoria), ’Monotoca’ (18; southwestern Western Australia, southeastern South Australia to eastern Queensland, Tasmania; non-monophyletic), Planocarpa (3; P. nitida, P. petiolaris, P. sulcata; Tasmania), Styphelia (15; southwestern Western Australia, southeastern South Australia to southeastern Queensland, Tasmania), Trochocarpa (12; Malesia to New Guinea, northeastern Queensland, eastern New South Wales, eastern Victoria, Tasmania). – Chile, southern Indochina, Malesia to New Guinea, Australia, Tasmania, Melanesia, New Zealand, the Hawaiian Islands and other islands in the Pacific. Ericoid mycorrhiza? Nodes in Richeeae ≥3:≥3. Leaves xeromorphic, ericoid, not recurved, often with apical spine, in Cosmelieae and Richeeae with sheathing base. Venation parallel or palmate; mid-vein usually indistinct (in Prionoteae distinct, with three veins from leaf base). Foliar vascular bundles with well developed abaxial fibrous envelopes. Adaxial cap absent. Leaf epidermis lignified (also in Lyonieae in Vaccinioideae); epidermal cells rectangular, arranged in longitudinally parallel rows; anticlinal epidermal cells walls sinuous (at least on abaxial side of leaf); abaxial epidermis and associated hypodermis detach from mesophyll in Archerieae. Stomata in Richeeae brachyparacytic; in Styphelieae parallel to foliar longitudinal axis. Abaxial foliar surface in Oligarrheneae without ribbon wax and papillae; adaxial cuticular wax crystalloids absent in Richeeae. Multicellular hairs and leaf teeth usually absent (present in Prionoteae). Inflorescence terminal or axillary, usually spicate; flowers often solitary axillary with numerous floral prophylls (absent in Richeeae). Inflorescence bracts membranous. Petals in Oligarrheneae with valvate or induplicate valvate aestivation. Stamens usually as many as sepals, antesepalous (in Oligarrheneae two to five). Filaments usually adnate to corolla tube (epipetalous). Anthers disporangiate (monothecal), longicidal (usually dehiscing by a single slit, in Prionoteae by two slits), without appendages. Tapetum with uninucleate cells. Pollen grains in Styphelieae shed as pseudomonads (only one out of four meiospores developing). Pistil in Oligarrheneae composed of two carpels. Ovule in Oligarrheneae one per carpel. Ovules usually epitropous (in Lysinema in Epacrideae apotropous). Fruit in Oligarrheneae a nut, in Styphelieae a drupe; corolla persistent in fruit. – A probable topology is (Crayn & Quinn 2000): [Prionoteae+[Archerieae+[Oligarrheneae+[Cosmelieae+[Richeeae+[Epacrideae+Styphelieae]]]]]].

Cladogram (simplified) of Vaccinioideae based on morphology and DNA sequence data (Kron, Judd & Crayn 1999).

Vaccinioideae Arn., Botany: 118. 9 Mar 1832 [‘Vaccinieae’] (under construction)

45/1.450–1.480. Vaccinieae Rchb., Fl. Germ. Excurs. 1(3): 203. Jul-Dec 1831. ’Vaccinium’ (c 150?; temperate and arctic-alpine regions on the Northern Hemisphere, mountains in Central and southeastern Africa and Madagascar, the Hawaiian Islands, mountains in tropical America; polyphyletic), Gaylussacia (>50; eastern Canada, eastern United States, the Andes in South America; incl. Vaccinium pro parte), Oxycoccus (4; O. erythrocarpus, O. macrocarpus, O. microcarpus, O. palustris; temperate and arctic-alpine regions on the Northern Hemisphere), Symphysia (15; Central America, the West Indies), Agapetes (c 400; tropical Asia from India to New Guinea; incl. Vaccinium pro parte), Didonica (4; D. crassiflora, D. panamensis, D. pendula, D. subsessilis; Costa Rica, Panamá), Notopora (5; N. auyantepuiensis, N. cardonae, N. chimantensis, N. schomburgkii, N. smithiana; eastern Venezuela), Orthaea (34; Mexico, Central America, the Andes south to Bolivia), Gonocalyx (9; Central America), Dimorphanthera (75–80; Malesia, with their highest diversity on New Guinea), Paphia (c 20; eastern New Guinea, eastern Queensland, New Caledonia, Fiji; incl. Dimorphanthera keysseri), Cavendishia (100–110; southern Mexico, Central America, tropical South America; incl. Thibaudia pro parte), Macleania (35–40; southern Mexico, Central America, western tropical America), Psammisia (c 60; tropical America), Satyria (c 25; southern Mexico, Central America, tropical South America), Mycerinus (3; M. chimantensis, M. sclerophyllus, M. viridiflorus; the Guayana Highlands), Polyclita (1; P. turbinata; Bolivia), Anthopteropsis (1; A. insignis; central Panamá), Ceratostema (23–32; mountain regions in South America, with their highest diversity in the Andes in Ecuador), Semiramisia (3; S. karsteniana, S. pulcherrima, S. speciosa; northern Andes), Oreanthes (7; O. buxifolius, O. ecuadorensis, O. fragilis, O. glanduliferus, O. hypogaeus, O. rotundifolius, O. sperlingii; the Andes in Ecuador), Siphonandra (5; S. boliviana, S. elliptica, S. magnifica, S. nervosa, S. santa-barbarensis; the Andes), Pellegrinia (6; P. coccinea, P. dichogama, P. grandiflora, P. guascensis, P. harmsiana, P. hirsuta; the Andes), ’Disterigma’ (c 35; northern and central Andes; polyphyletic), Utleya (1; U. costaricensis; Costa Rica), Sphyrospermum (>22; mountains in tropical America; incl. Disterigma pro parte), Rusbya (1; R. taxifolia; northern Bolivia), Themistoclesia (c 25; northern Andes), Plutarchia (11; Colombia, Ecuador), Anthopterus (12; the Andes; incl. Thibaudia pro parte), ’Thibaudia’ (c 60; Central America, northwestern tropical South America; polyphyletic), Demosthenesia (11; the Andes), ’Diogenesia’ (13; the Andes; polyphyletic). – Andromedeae Klotzsch, Pfl.-Abbild. Beschr.: 16. 1838. Andromeda (2; A. glaucophylla, A. polifolia; temperate regions on the Northern Hemisphere), Zenobia (1; Z. pulverulenta; southeastern United States). – Gaultherieae Nied. in Engl. Bot. Jahrb. Syst. 11: 145. 18 Jun 1889. Chamaedaphne (1; C. calyculata; temperate regions on the Northern Hemisphere), Eubotrys (2; E. racemosa, E. recurva; southeastern United States), Gaultheria (c 135; East Asia to Malesia, Australia, New Zealand, Alaska, Canada, United States, Mexico, Central America, South America), ’Leucothoe’ (5; L. axillaris, L. davisiae, L. fontanesiana, L. griffithiana, L. keiskei; the Himalayas, Japan, North America; polyphyletic), Tepuia (7; T. cardonae, T. intermedia, T. multiglandulosa, T. speciosa, T. tatei, T. vareschii, T. venusta; the Guayana Highlands). – Lyonieae Kron et Judd in K. A. Kron et al., Amer. J. Bot. 86: 1298. 15 Sep 1999. Agarista (c 30; tropical Africa, Madagascar, the Mascarene Islands, southeastern United States, Mexico, Central America, tropical South America), Craibiodendron (5; C. henryi, C. scleranthum, C. stellatum, C. vietnamense, C. yunnanense; southern China, Southeast Asia), Lyonia (c 35; East and Southeast Asia, eastern United States, Mexico, the West Indies), Pieris (7; P. cubensis, P. floribunda, P. formosa, P. japonica, P. nana, P. phillyreifolia, P. swinhoei; the Himalayas to East Asia and eastern Siberia, southeastern United States, Cuba). – Oxydendreae Cox ex Reveal in Phytoneuron 2012-37: 218. 23 Apr 2012.Oxydendrum (1; O. arboreum; eastern United States). – Cosmopolitan. Often epiphytic. Mycorrhiza ericoid. Phellogen in ‘Vaccinium’ and Agapetes superficial. Secondary phloem in Lyonieae often with bands of fibres. Apical bud aborted. Epidermis in Lyonieae lignified. Stomata often paracytic (in Lyonieae often anomocytic). Inflorescence usually axillary. Pedicel often articulated. Hypogyny frequent. Anthers spurred, with two or four bristles or without appendages; abaxial side of anthers with disintegration tissue. Tapetum cells in Vaccinium binucleate. Ovary often inferior, in ‘Vaccinium’ and Agapetes appearing decemlocular. Stigma truncate. x = 12.

Cladogram of Vaccinieae based on DNA sequence data (Kron, Powell & Luteyn 2002). – Agapetes clade A: Vaccinium caudatifolium, V. filiforme, V. lancifolium. – Agapetes clade B: Agapetes, V. gaultheriifolium, V. nummularia. – Bracteata-Oarianthe clade: V. cercidifolium, V. leptospermoides, V. summifaucis, V. finisterrae, V. horizontale. – Myrtillus clade: Costera endertii, V. hirtum, V. smallii, V. parvifolium, V. myrtillus, V. dentatum, V. reticulatum. – Orthaea-Notopera clade: V. crenatum, Notopora schomburgkii, Orthaea apophysata, Orthaea venamensis. – Vaccinium clade: V. macrocarpon, V. vitis-idaea. – East Malesian clade: Paphia, Dimorphanthera. – Andean clade A: Thibaudia floribunda, Anthopterus, Themistoclesia, Diogenesia, Demosthenesia; Satyria, Orthaea fimbriata, Cavendishia (incl. Thibaudia jahnii). – Andean clade B: Ceratostema, Oreanthes, Sphyrospermum (incl. Disterigma pro parte), Macleania, Psammisia, Plutarchia, Polyclita (incl. Thibaudia macrocalyx), Satyria boliviana, Thibaudia densiflora, Semiramisia, Thibaudia parvifolia, Siphonandra, Thibaudia pachyantha. – Meso-American/Caribbean clade: Symphysia, Disterigma trimerum, Utleya, V. poasanum, Gonocalyx, Macleania megabracteolata. – Vaccinium Section Hemimyrtillus may be sister-group to the remaining Vaccinieae, yet with weak support (Powell & Kron 2002).

Fouquieriales DC. in C. F. P. von Martius, Consp. Regn. Veg.: 50. Sep-Oct 1835 [’Fouquieriaceae’]; Fouquierineae Engl., Syllabus, ed. 2: 153. Mai 1898

Habit Bisexual, deciduous shrubs or small trees. Xerophytic, often stem succulents, spiny, with little branched long and short shoots. Periderm transparent, often exfoliating.

Vegetative anatomy Phellogen ab initio superficially (in root) or deeply seated (often outer-cortical). Layers of fibre cells alternating with layers of cork cells in stem phelloderm. Stem cortex reticulum consisting of groups of superficial sclereids and assimilating tissue, and inside this partly water-storing and partly starch-storing tissue. Medulla septated by diaphragms. Vessel elements with usually simple (rarely scalariform) perforation plates; lateral pits alternate or opposite, simple and/or bordered pits. Imperforate tracheary xylem elements fibre tracheids with bordered pits, non-septate (also vasicentric tracheids). Wood rays multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty. Xylem parenchyma well developed and water-storing. Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace. Prismatic calciumoxalate crystals abundant.

Leaves Alternate (spiral), simple, entire, heteromorphic, with ? ptyxis. Stipules and leaf sheath absent. Petioles of long shoots modified into long cortical spines, with base proceeding down stem as coarse ridges. Axillary short shoots with normal leaves. Petiole vascular bundles? Venation pinnate. Stomata anomocytic. Cuticular wax crystalloids absent. Epidermis with mucilage cells (with mucilaginous inner walls). Leaf margin entire.

Inflorescence Terminal or axillary, spike- or raceme-like, panicle or corymbo-paniculate.

Flowers Actinomorphic. Hypogyny. Sepals five, with imbricate quincuncial aestivation, spiral, free, with membranous margin, persistent. Petals five, with imbricate quincuncial aestivation, connate into tube. Base of ovary nectariferous. Disc small, annular.

Androecium Stamens usually ten (rarely up to 23), diplostemonous (antepetalous stamens sometimes duplicated). Filaments free, free from or slightly adnate to corolla tube, sometimes with ligulate spur at base. Anthers dorsifixed, versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolporate, shed as monads, bicellular at dispersal. Exine semitectate, with columellate infratectum, reticulate, often striate.

Gynoecium Pistil composed of three connate carpels. Ovary superior, unilocular, with nectariferous vascularized tissue at base. Stylodia three, connate in lower part. Stigmas punctate, type? Pistillodium absent.

Ovules Placentation at anthesis parietal with deeply intrusive placentae (axile at base), finally free central. Ovules six to c. 20 per carpel, anatropous, ascending, apotropous, bitegmic, tenuinucellar. Micropyle endostomal. Outer integument three or four cell layers thick. Inner integument (four to) seven or eight cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development cellular. Endosperm haustoria chalazal and micropylar. Embryogenesis asterad.

Seeds Aril? Seed coat winged. Testa multiplicative, gradually crushed; hypodermis with band-shaped thickenings. Tegmen multiplicative, crushed. Perisperm not developed. Endosperm thin, oily and proteinaceous, or absent. Embryo small, straight, well differentiated, chlorophyll? Cotyledons two, flat. Germination phanerocotylar.

Phytochemistry Flavonols, cyanidin, Route I secoiridoids, dammarane, ellagic acid, pelargonidin (in red-flowered species), caffeic acid, indole alkaloids and their glycosides, and saponins present. Myricetin and cyanogenic compounds not found.

Richard in J. B. G. M. Bory de Saint-Vincent, Dict. Class. Hist. Nat. 9: 269. 25 Feb 1825 [‘Lecythideae’], nom. cons.

Belvisiaceae R. Br. in Trans. Linn. Soc. London 13: 222. Mai-Jun 1821 [‘Belviseae’], nom. illeg.; Napoleonaeaceae A. Rich. in J. B. G. M. Bory de Saint-Vincent, Dict. Class. Hist. Nat. 11: 432. 10 Feb 1827 [‘Napoleoneae’]; Barringtoniaceae (DC.) F. Rudolphi, Syst. Orb. Veg.: 56. 5-12 Jul 1830 [‘Barringtonieae’], nom. cons.; Barringtoniales DC. in C. F. P. von Martius, Consp. Regn. Veg.: 65. Sep-Oct 1835 [‘Barringtonieae’]; Belvisiales R. Br. in C. F. P. von Martius, Consp. Regn. Veg.: 63. Sep-Oct 1835 [’Belvisieae’], nom. illeg.; Gustaviaceae Burnett, Outl. Bot.: 717, 1092, 1135. Feb 1835; Lecythidales Poit. in C. F. P. von Martius, Consp. Regn. Veg.: 65. Sep-Oct 1835 [’Lecythideae’]; Barringtoniineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 583, 602. 1846; Scytopetalaceae Engl. in Engler et Prantl, Nat. Pflanzenfam. Nachtr. 1: 242. Oct 1897, nom. cons.; Scytopetalineae Engl., Syllabus, ed. 2: 150. Mai 1898; Rhaptopetalaceae Tiegh. ex Solereder, Syst. Anat. Dicot. Ergänz.: 53. Mai 1908; Asteranthaceae R. Knuth in Engler, Pflanzenr., IV, 219b (Heft 105): 1. 22 Aug 1939, nom. cons.; Foetidiaceae (Engl.) Airy Shaw in Kew Bull. 18: 258. 8 Dec 1965; Lecythidanae Takht. ex Reveal in Novon 2: 236. 13 Oct 1992

Fossils The fossil flower Lecythidoanthus kugleri has been described from the Miocene of Trinidad. Fossil wood, fruits and pollen grains assigned to Lecythidaceae were recorded from Late Cretaceous (Maastrichtian) and Cenozoic sediments in India and Europe.

Habit Bisexual, usually evergreen trees (sometimes shrubs or lianas; in Scytopetaloideae rarely perennial herbs). Foetidia has evil-smelling wood. Bark often fibrous.

Vegetative anatomy Phellogen ab initio superficial. Stem cortex with vascular bundles. Primary medullary rays narrow or wide. Vessel elements usually with simple (sometimes scalariform, rarely reticulate) perforation plates; lateral pits alternate or opposite, simple and/or bordered pits. Imperforate tracheary xylem elements tracheids (in Allantoma), fibre tracheids or libriform fibres usually with simple (sometimes bordered) pits, septate or non-septate (also vasicentric tracheids). Wood rays uniseriate to multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty vasicentric, reticulate or banded (rarely aliform, confluent or scalariform). Tyloses often abundant (sometimes sclerotic). Secondary phloem stratified into hard fibrous and soft parenchymatous layers, often with unilaterally thickened cells with crystals (sometimes with cuneate rays). Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace, or ≥3:≥3, trilacunar to multilacunar with three or more traces. Cristarque cells sometimes present. Cells with calciumoxalate crystals or silica bodies. Prismatic crystals abundant.

Trichomes Hairs unicellular or multicellular, usually uniseriate (sometimes stellate) or absent; glandular hairs sometimes present.

Leaves Usually alternate (sometimes distichous; rarely opposite), simple, entire, often coriaceous, with supervolute or involute ptyxis. Stipules usually absent (in Asteranthos and Scytopetalum small, paired, cauline, caducous); leaf sheath absent. Colleters present. Petiole vascular bundle transection arcuate or annular; petiole with numerous vascular bundles. Venation pinnate, sometimes brochidodromous. Stomata usually anisocytic (sometimes paracytic or anomocytic). Cuticular wax crystalloids as polygonal platelets. Domatia as pits (Combretodendron) or absent. Secretory cavities absent. Mesophyll in Napoleonaeoideae with idioblasts containing ethereal oils and sclerenchymatous idioblasts containing prismatic calciumoxalate crystals. Leaf margin usually entire (sometimes serrate or crenate).

Inflorescence Terminal or axillary, spike- or raceme-like, panicle, fasciculate, thyrsoid, botryoid cymose, or racemose, or flowers solitary axillary (each flower in Napoleonaeoideae subtended by four to seven often glanduliferous bracts, extrafloral nectaries).

Flowers Actinomorphic or zygomorphic (especially androecium), often large. Pedicel articulated. Hypanthium present or absent. Epigyny or half epigyny (in Scytopetaloideae usually hypogyny). Sepals (two or) four to six (to twelve), usually with imbricate (sometimes valvate) aestivation, persistent or caducous (in Asteranthos accrescent in fruit), usually more or less connate. Petals three to six (to eight, ten, twelve, 16, or 18), with imbricate aestivation, caducous, usually connate (absent in Foetidioideae and probably in Napoleonaeoideae and Scytopetaloideae). Nectaries often present at base of staminodia on androphore. Nectariferous disc annular, intrastaminal or extrastaminal, present in Napoleonaeoideae, Barringtonioideae and Foetidia (in Asteranthos formed by fused staminodia).

Androecium Stamens ten to more than 1.200, with various positions, often whorled, usually centrifugally (sometimes centripetally) developing from annular primordium, often (some Lecythidoideae) on annular or asymmetrical androphore (consisting of staminodia and/or fertile stamens), sometimes curved and hood-like above ovary. Filaments usually more or less connate (in Foetidioideae and Napoleonaeoideae free), free from tepals (in Napoleonaeoideae and Scytopetaloideae adnate to base of pseudocorolla probably consisting of staminodial tissue). Anthers basifixed, often versatile, usually tetrasporangiate (in Napoleonaeoideae disporangiate), extrorse, latrorse or introrse, usually longicidal (dehiscing by longitudinal slits; in Scytopetaloideae sometimes poricidal, with apical short slits or pores). Tapetum secretory or amoeboid-periplasmodial. Staminodia (sometimes with fodder pollen in anthers) often several to numerous on androphore or pseudocorolla, extrastaminal (outer staminodial whorl in Napoleonaea forming pseudocorolla, and two inner staminodial whorls forming corona; staminodial whorl in Scytopetaloideae forming pseudocorolla).

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolpate or tricolpor(oid)ate or syntricolpate (Barringtonioideae), shed as monads, bicellular or tricellular at dispersal. Exine tectate, with granular infratectum, smooth, perforate to microreticulate, foveolate or scabrate-verrucate.

Gynoecium Pistil composed of two to six (to eight) connate usually antesepalous (sometimes antepetalous) carpels. Ovary inferior to superior, unilocular to sexalocular (to octalocular). Style single, simple, or absent. Stigma one, capitate, punctate or trilobate to octalobate, papillate or non-papillate, Dry or Wet type. Pistillodium absent.

Ovules Placentation apical, apical-axile to basal-axile (in Eschweilera basal; in Foetidia axile, with peltate placentae). Ovules (one or) two to c. 115 per carpel, usually anatropous (rarely campylotropous), pendulous, horizontal or ascending, bitegmic, tenuinucellar. Micropyle usually endostomal (in, e.g., Couroupita exostomal). Outer integument six to 18 cell layers thick. Inner integument three to seven cell layers thick. Archespore multicellular. Megagametophyte monosporous, Polygonum type. Antipodal cells ephemeral. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit A drupe (in Foetidioideae and Barringtonioideae, sometimes winged, in Napoleonaeoideae with persistent calyx and style), large to very large, indehiscent and juicy inside, or a woody pyxidium (rarely a loculicidal capsule or a berry-like fruit).

Seeds Aril present in many species of Lecythis. Seed coat mesotestal, in Cariniana and Couroupita winged. Testa vascularized, multiplicative, often strongly lignified. Exotestal cells of various thickness, palisade, or with sinuous anticlinal walls. Mesotesta often sclerotized. Endotesta? Tegmen? Perisperm not developed. Endosperm usually very sparse or absent (in Scytopetaloideae copious, with amyloidiferous cell walls and usually ruminate). Embryo large, straight or curved, undifferentiated or well differentiated, chlorophyll? Cotyledons two, fleshy planoconvex or foliaceous, or undifferentiated. Hypocotyl often strongly developed. Germination phanerocotylar or cryptocotylar.

Cytology n = 13 (26) (Barringtonioideae); n = 16 (Napoleonaeoideae); n = 11, 18, 21 (Scytopetaloideae); n = 17 (18) (34) (Lecythidoideae); n = ? (Foetidioideae)

DNA Mitochondrial intron coxII.i3 lost (Bertholletia). Mitochondrial coxI intron present (Barringtonia).

Phytochemistry Flavonols (quercetin etc.), cyanidin, oleanolic acid derivatives, ellagic and gallic acids, methylated ellagic acids, proanthocyanidins (prodelphinidins), triterpene saponins, and cyanogenic compounds present. Kaempferol and alkaloids not found. Selenium accumulated in some species. Aluminium accumulated in some Scytopetaloideae.

Use Ornamental plants, edible seeds (Bertholletia excelsa, Lecythis zabucajo), timber.

Systematics The sister-group relationships of Lecythidaceae are unresolved. They are part of a trichotomy also comprising [Fouquieriaceae+Polemoniaceae] and the majority of Ericales.

Lecythidaceae and Sapotaceae have seeds with multiplicative lignified testa.

A plausible topology of the Lecythidaceae subclades is the following: [Napoleonaeoideae+[Scytopetaloideae+[Lecythidoideae+[Barringtonioideae+Foetidioideae]]]].

Napoleonaeoideae (R. Br.) Benth. in W. J. Hooker, Niger Fl.: 360. Nov-Dec 1849 [‘Napoleoneae’]

2/c 13. Napoleonaea (c 10; tropical West and Central Africa to Angola and Zambia), Crateranthus (3; C. congolensis, C. le-testui, C. talbotii; southern Nigeria, Gabon, Congo). – Tropical West and Central Africa. Secondary xylem containing crystal chains. Leaves distichous, with supervolute ptyxis. Stipules present or absent. Lamina often with glands on abaxial side at base. Leaf margin serrate. Sepals three (Crateranthus) or five (Napoleonaea). Petals with valvate aestivation, connate, plicate. Nectaries present on abaxial side of sepals. Filaments adnate to corolla. Anthers long (in Napoleonaea ten, pairwise, antepetalous, extrorse, monothecal). Corona formed by two inner staminodial whorls. Carpels antepetalous, initiated prior to stamens. Style absent. Stigma wide, quinquangular, flat. Ovules four per carpel, in Napoleonaea collateral in pairs. Endothelium absent. Fruit a drupe with one or several seeds. Embryo curved. n = 16. – Napoleonaeoideae have mixed simple and scalariform vessel perforation plates, scalariform vessel-ray pitting, and high multiseriate wood rays, features also present in Scytopetaloideae. The wood structure of Napoleonaea is distinct, but its hypothetical sister-group Crateranthus strongly resembles Scytopetaloideae.

[Scytopetaloideae+[Lecythidoideae+[Barringtonioideae+Foetidioideae]]]

Scytopetaloideae O. Appel in Bot. J. Linn. Soc. 121: 225. 17 Sep 1996

6/22–23. Asteranthos (1; A. brasiliensis; Venezuela, northern Brazil), Oubanguia (3; O. africana, O. alata, O. laurifolia; tropical West and Central Africa), Scytopetalum (3; S. klaineanum, S. pierreanum, S. tieghemii; tropical West and Central Africa), Rhaptopetalum (11–12; tropical West and Central Africa), Pierrina (1; P. zenkeri; Cameroon, Gabon, Equatorial Guinea), Brazzeia (3; B. congoensis, B. longipedicellata, B. soyauxii; Central Africa). – Tropical West and Central Africa, northern Brazil. Leaf traces passing two internodes prior to entering leaves. Sclereids present. Leaves distichous. Stipules minute. Pedicel articulated. Usually hypogyny (sometimes half hypogyny). Sepals connate. Petals with valvate aestivation, calyptrate, caducous, sometimes connate at base (sometimes absent). Nectariferous disc usually present (nectary sometimes absent). Filaments connate at base, adnate to corolla and to staminodia. Tapetum secretory. Staminodia six to 28, petalous, connate. Pollen grains tricolpate or sometimes tricolporoidate. Staminodia six to 16 or 24 to 28 (Asteranthos), connate, petaloid. Style long, thin. Stigma punctate. Ovules two or four per carpel. Micropyle prolonged. Endothelium present. Fruit a capsule or indehiscent, often one-seeded. Seed coat hairy, usually ruminate (not in Oubanguia). Endosperm with hemicellulose. Embryo J-shaped. n = 11, 18, 21. Aluminium accumulated in some species. – Asteranthos resembles Napoleonaeoideae in its morphology and phytochemistry, but is placed in Scytopetaloideae by DNA sequence analyses.

[Lecythidoideae+[Barringtonioideae+Foetidioideae]]

Embryo usually with starch.

Lecythidoideae Beilschm. in Flora 16(Beibl. 7): 98, 108. 14 Jun 1833 [‘Lecythideae’]

10/c 220. Gustavia (c 40; Central America, tropical South America), Grias (12; Panamá to Peru), Allantoma (8; northeastern tropical South America), Cariniana (9; tropical South America), Couroupita (3; C. guianensis, C. nicaraguarensis, C. subsessilis; Central America, tropical South America), Corythophora (4; C. alta, C. amapaensis, C. labriculata, C. rimosa; French Guiana, Suriname, Brazil), Bertholletia (1; B. excelsa; tropical South America), Couratari (c 20; tropical South America), ’Eschweilera’ (c 95; southern Mexico, Central America, tropical South America; non-monophyletic), ’Lecythis’ (27; Central America, tropical South America; non-monophyletic). – Southern Mexico, Central America, northern and central South America to Uruguay. Bark fibrous. Cortical vascular bundles in Gustavia inverted. Secondary xylem with crystal chains. Leaves spiral or distichous, with involute ptyxis. Extraflora nectaries present in Gustavia. Flowers usually (secondarily?) zygomorphic (sometimes actinomorphic). Nectariferous disc present or absent. Filaments contracted at apex, connate at base forming basal ring. Pollen grains tricolp(oroid)ate, sometimes heteromorphic. Fodder pollen produced in many species. Style usually short (sometimes long). Endothelium absent. Fruit usually a pyxidium (rarely indehiscent). Wing-shaped aril or swollen funicle present or absent. Embryo often curved, with hypocotyl or long radicula and foliaceous (sometimes folded) or thick cotyledons. n = 17 (18) (34).

[Barringtonioideae+Foetidioideae]

Cortical vascular bundles inverted. Leaves with supervolute ptyxis. Fruit indehiscent.

Barringtonioideae Beilschm. in Flora 16(Beibl. 7): 98, 104. 14 Jun 1833 [‘Barringtonieae’]

5/c 84. Petersianthus (2; P. macrocarpus: tropical West and Central Africa to Angola and Congo; P. quadrialatus: the Philippines), Barringtonia (c 70; East Africa, Madagascar, tropical Asia to the Ryukyu Islands and Micronesia), Chydenanthus (1; C. excelsus; Burma, the Andaman and Nicobar Islands, Malesia to New Guinea), Abdulmajidia (2; the Malay Peninsula), Careya (3; C. arborea: Afghanistan to the Malay Peninsula; C. herbacea: the Himalayas; C. valida: the Andaman Islands), Planchonia (8; the Andaman and Nicobar Islands, the Malay Peninsula to New Guinea, Solomon Islands and tropical Australia). – Tropical East Africa, Madagascar, South and Southeast Asia, Malesia to New Guinea, tropical Australia, Melanesia to the Ryukyu Islands and Micronesia. Secondary xylem without crystal chains. Nodes 1:1. Leaves spiral. Glands often replacing stipules. Nectariferous disc annular. Filaments connate at base forming basal ring. Pollen grains syn(tri)colpate, with strong margin ridge at colpus. Style long. Endothelium usually absent. Fruit usually one-seeded. Embryo with hypocotyl or long radicula and foliaceous cotyledons. n = 13 (26). – Barringtonioideae and Lecythidoideae both have vessel elements with exclusively simple perforations and two types of vessel ray pitting. However, they may be distinguished from each other by the size of intervessel pitting, the shape of body ray cells arranged in multiseriate rays, and the type of crystalliferous axial parenchyma cells.

Foetidioideae Engl., Syllabus, ed. 1: 146. Apr 1892

1/18. Foetidia (18; Madagascar, the Comoros, the Mascarene Islands, one species, F. africana, in tropical East Africa). – Secondary xylem with crystal chains. Nodes 1:1. Leaves elongating in bud. Sepals with valvate aestivation, lignified. Petals absent. Nectariferous disc indistinct. Filaments free. Anthers introrse. Style trilobate or quadrilobate. Placentae peltate, with ovules inserted in two rows. Endothelium present. n = ? – The isolated position of Foetidia (Foetidioideae) may be supported by a unique type of vessel ray pitting similar in shape and size to intervessel pitting (distinctly bordered, <5 mm).

Cladogram of Lecythidaceae based on morphology, anatomy and DNA sequence data (Morton, Mori & al. 1997).

Cladogram of Lecythidaceae based on DNA sequence data (Mori & al. 2007).

Distribution Tropical Africa, Southeast Africa, Madagascar, Yemen, Sri Lanka, East Asia to Japan, Southeast Asia, Malesia to New Guinea, Melanesia, Queensland, islands in the Pacific.

Habit Usually bisexual (often functionally polygamous, or functionally or cryptically dioecious?), evergreen trees, shrubs or lianas.

Vegetative anatomy Phellogen ab initio superficial? Vessel arranged in radial multiples. Vessel elements with simple and/or scalariform (sometimes reticulate) perforation plates; lateral pits alternate, bordered pits. Imperforate tracheary xylem elements libriform fibres? with simple pits, septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse, or paratracheal scanty. Sieve tube plastids S type? Nodes unilacunar? with ? leaf trace; lateral vascular bundles arising approx. halfway down internode below supported leaf. Schizogenous secretory canals with tannins etc. Parenchyma cells with calciumoxalate druses.

Leaves Alternate (spiral or distichous), simple, entire, with induplicate ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection annular. Venation pinnate, often indistinct. Stomata anisocytic? Cuticular wax crystalloids? Lamina with well developed schizogenous secretory canals. Abaxial epidermis often with groups of prismatic calciumoxalate druses. Leaf margin serrate or entire.

Inflorescence Axillary simple or compound raceme. Floral prophylles (bracteoles) two.

Flowers Actinomorphic. Half epigyny (perigyny). Sepals four or five (or six), with quincuncial aestivation, persistent, free or connate. Petals four or five (or six), with induplicate-valvate aestivation, connate into campanulate corolla. Petals and stamens developing from common primordia; stamen primordium smaller than petal primordium. Tepals with well developed and distinct canals. Nectaries present on ovary wall. Disc absent.

Androecium Stamens usually four or five (sometimes six), haplostemonous, antepetalous, alternisepalous. Filaments connate at base, adnate to corolla tube (epipetalous). Anthers dorsifixed, versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Female flowers with staminodia?

Pollen grains Microsporogenesis simultaneous? Pollen grains colpate or colpor(oid)ate, shed as monads, bicellular at dispersal. Exine semitectate, with columellate infratectum, subreticulate.

Gynoecium Pistil composed of usually three or four connate antepetalous carpels. Ovary semi-inferior, unilocular, nectariferous. Style single, simple (hollow?). Stigma truncate, capitate or bilobate to quinquelobate, type? Male flowers with pistillodium?

Ovules Placentation free central. Ovules numerous per ovary, anatropous, apotropous, bitegmic, tenuinucellar. Micropyle endostomal? Outer integument approx. two cell layers thick. Inner integument three or four cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis onagrad.

Fruit A many-seeded drupaceous berry with persistent calyx. Mesocarp often thin. Endocarp somewhat lignified.

Seeds Aril absent. Exotesta? Endotesta with rhomboidal crystals. Tegmen? Perisperm not developed. Endosperm copious, amyloidiferous and lipidiferous. Embryo small, straight, chlorophyll? Cotyledons two, small. Germination?

Phytochemistry Insufficiently known. Flavonols (kaempferol, quercetin, myricetin?), alkaloids, triterpene saponins, and maesaquinone and other quinones present. Cyanidin? Ellagic acid? Proanthocyanidins? Glycosides? Cyanogenic compounds?

Use Medicinal plants, seed oil, spices (Maesa indica), fish poisons, insecticides, timber.

Berchtold et Presl., Přir. Rostlin: 218. Jan-Apr 1820 [’Marcgraviae’], nom. cons.

Marcgraviales Juss. in C. F. P. von Martius, Consp. Regn. Veg.: 61. Sep-Oct 1835 [’Marcgraviaceae’]; Noranteaceae (Choisy) DC. ex Mart., Consp. Regn. Veg.: 61. Sep-Oct 1835

Distribution Southern Mexico, Central America, the West Indies, northern to central South America.

Fossils Fossil pollen sometimes attributed to Marcgravia and Norantea s.lat. have been reported from Early Oligocene sediments in Puerto Rico. Fossil wood from Cenozoic layers on Sumatra has been described under the name of Ruyschioxylon.

Habit Bisexual evergreen shrubs, often epiphytic or semi-epiphytic, or lianas (rarely small trees).

Vegetative anatomy Phellogen ab initio superficial. Primary medullary rays alternating wide and narrow. Primary vascular tissue a cylinder, without separate vascular bundles. Vessel elements with simple and/or scalariform perforation plates; lateral pits alternate or opposite, simple or bordered pits. Imperforate tracheary xylem elements fibre tracheids or libriform fibres with simple and/or bordered pits, usually septate. Wood rays uniseriate to multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty vasicentric. Sieve tube plastids Ss type. Nodes 1:1, unilacunar with one leaf trace. Cortical parenchyma with idioblasts containing sclereids or calciumoxalate raphides (in raphid sacs).

Trichomes Hairs usually unicellular (rarely multicellular); glands frequent.

Leaves Alternate (spiral or in Marcgravia distichous), simple, entire, often coriaceous, with supervolute ptyxis; in Marcgravia often heterophyllous, with small cordate leaves on sterile shoots with climbing roots and ordinary leaves on fertile root-less shoots. Stipules and leaf sheath absent. Petiole vascular bundles? Extrafloral nectaries often present on basal parts of abaxial surface of lamina. Venation pinnate, brochidodromous, indistinct. Stomata staurocytic. Cuticular wax crystalloids as platelets. Mesophyll with sclerenchymatous idioblasts containing calciumoxalate raphides. Abaxial side of lamina often with nectariferous glands (in Marcgravia also along margin). Domatia as pits. Leaf margin usually entire (sometimes serrate). Leaf apex with often caducous mucro.

Inflorescence Terminal raceme, umbel or spike. Flower in some species with petaloid bract adnate to pedicel and modified into leaf-, spur-, sac-, cup- or pitcher-like extrafloral nectary (nectaries in Marcgravia inserted on central aborted flowers).

Flowers Actinomorphic. Hypogyny. Sepals usually five (in Marcgravia four), with imbricate quincuncial aestivation, unequal in size, persistent, free or somewhat connate at base. Petals three to five, with imbricate aestivation, free or more or less connate (in Marcgravia four, in the form of a calyptra falling off at beginning of anthesis). Nectariferous disc absent.

Androecium Stamens (three to) five to more than 100, in one or two whorls. Filaments filiform to flattened, free or connate at base, usually free from tepals (rarely adnate to petal bases). Anthers basifixed or subbasifixed, non-versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, with uninucleate cells. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolporate (sometimes tetracolporate), shed as monads, bicellular at dispersal. Exine tectate to semitectate, with usually columellate infratectum, imperforate, perforate or reticulate.

Gynoecium Pistil composed of two to eight connate carpels. Ovary superior, unilocular at first, gradually entirely or incompletely bilocular to 15-(to 20-)locular. Style single, simple, short, or absent. Stigma simple or lobed, Dry type. Pistillodium absent.

Ovules Placentation strongly intrusively parietal (seemingly axile). Ovules five to c. 20 (in Marcgraviastrum much more than 20) per carpel, anatropous, (incompletely) bitegmic, incompletely tenuinucellar. Micropyle endostomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development cellular. Endosperm haustorium micropylar. Embryogenesis?

Fruit A loculicidal and septifragal capsule (or irregularly dehiscing) with persistent calyx, style and stigma. Pericarp leathery. Placentae carnose.

Seeds Aril often present. Seed coat exotestal? Exotestal cells enlarged, with inner walls strongly thickened, with theoid exotestal thickenings. Endotesta? Tegmen? Perisperm not developed. Endosperm sparse, starchy, or absent. Embryo large, straight or somewhat curved, well differentiated, chlorophyll? Cotyledons two, small to large. Germination phanerocotylar.

Phytochemistry Flavonols (myricetin), terpenes, tannins, proanthocyanidins, leucopelargonidin, alkaloids, and triterpene saponins present. Iridoids, ellagic acid and cyanogenic compounds not found. Carbohydrates sometimes stored as oligosaccharides with kestose or isokestose bonds (inulin).

Distribution Northeastern India, Japan, the Ryukyu Islands, Taiwan, Indochina, Malesia to New Guinea, southern Mexico, Guatemala, northwestern Colombia.

Habit Bisexual, achlorophyllous, herbaceous endophytes without rhizome or normal roots. Holoendoparasites on roots of Fagaceae (Castanopsis, Lithocarpus, Quercus, and Trigonobalanus).

Vegetative anatomy Mycorrhiza absent. Hypha-like threads of cells invading host plant and anastomosing forming reticulum in host root phloem. Phellogen absent. Secondary lateral growth absent. Vessels absent. Imperforate tracheary xylem elements? Wood rays absent. Axial parenchyma absent. Sieve tube plastids? Nodes absent. Crystals?

Leaves Opposite, reduced, scale-like (uppermost scales storing nectar), or absent. Stipules and leaf sheath absent. Venation? Stomata abnormal. Cuticular wax crystalloids cushion- or granule-shaped. Leaf margin entire.

Inflorescence Flowers terminal, solitary (flowers sometimes forming ‘fairy rings’ on ground).

Flowers Actinomorphic. Hypogyny. Tepals usually four (rarely five or six), with decussate-imbricate aestivation, somewhat petaloid, connate in lower part. Nectar-secreting stomata present on ovary and on base of androecium and upper scales. Disc?

Androecium Stamens numerous, entirely connate into cone-shaped tubular synandrous structure surrounding gynoecium, except for apical pore. Anthers arranged in several successive vertical whorls on different heights each whorl with approx. ten anthers, dorsifixed, non-versatile, polysporangiate (polythecate), extrorse, longicidal (anther whorls dehiscing circumscissely, splitting when pushed up by expanding style). Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains usually diporate or triporate (rarely tetraporate or possibly dicolpate?), shed as monads, bicellular at dispersal. Exine intectate?; ectexine reduced to small tubercles.

Gynoecium Pistil composed of eight to 15 connate carpels. Ovary superior, unilocular. Style single, simple, stout. Stigma hemispherical, sub-bilobate, type? Pistillodium absent.

Ovules Placentation parietal, with eight to 20 intrusive placentae. Ovules numerous per ovary, anatropous, unitegmic, tenuinucellar. Integument approx. two cell layers thick. Funicular obturator present. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio cellular. Endosperm haustoria? Embryogenesis solanad?

Fruit A many-seeded baccate pyxidium (dehiscing by horizontal slit along groove between style and ovary).

Seeds Aril absent. Funicle short, viscid. Seed coat endotestal. Exotestal cells with massive U-shaped thickenings. Endotesta? Perisperm not developed. Endosperm one-layered, oily, enclosing embryo. Embryo undifferentiated, quadricellular, without chlorophyll? Cotyledons two. Germination?

Lysimachiaceae Juss., Gen. Plant.: 95. 4 Aug 1789 [’Lysimachiae’]; Anagallidaceae Batsch ex Borkh., Bot. Wörterb. 1: 19. 1797 [’Anagallides’]; Ardisiaceae Juss. in Ann. Mus. Natl. Hist. Nat. 15: 350. 1810; Myrsinales R. Br. ex Bercht. et J. Presl, Přir. Rostlin: 251. Jan-Apr 1820 [‘Myrsineae’]; Myrsinopsida Bartl., Ord. Nat. Plant.: 121, 158. Sep 1830 [’Myrsineae’]; Aegicerataceae Blume, Nov. Plant. Expos.: 17. Aug-Dec 1833 [’Aegicereae’]; Lysimachiales Döll, Rhein. Fl.: 385. 24-27 Mai 1843 [‘Lysimachiae’]; Ardisiales J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 1025. 1846 [’Ardisiaceae’]; Coridaceae J. Agardh, Theoria Syst. Plant.: 332. Apr-Sep 1858 [’Corideae’]; Embeliaceae J. Agardh, Theoria Syst. Plant.: 140. Apr-Sep 1858 [’Embelieae’]

Habit Usually bisexual (rarely monoecious, polygamomonoecious or dioecious), evergreen or deciduous trees (in Aegiceras mangrove trees), shrubs, lianas or suffrutices, perennial or annual herbs. Cyclamen has a tuber formed by the hypocotyl.

Vegetative anatomy Phellogen ab initio superficially or deeply seated. Endodermis sometimes prominent? Vessel elements usually with simple (sometimes scalariform) perforation plates; lateral pits alternate, usually bordered pits. Imperforate tracheary xylem elements fibre tracheids or libriform fibres usually with simple pits, usually septate. Wood rays usually multiseriate, homocellular or heterocellular. Axial parenchyma paratracheal scanty vasicentric. Wood elements sometimes partially storied. Sieve tube plastids S type. Nodes usually 1:1, unilacunar with one leaf trace (sometimes 1:3 or 3:3, unilacunar or trilacunar with three traces). Secretory cells and schizogenous resinous canals with often reddish or blackish-red substance. Styloids, crystal sand and acicular calciumoxalate crystals present. Prismatic crystals often abundant.

Trichomes Hairs unicellular or multicellular, uniseriate or branched (sometimes articulated, stellate, dendritic, peltate or lepidote); glandular hairs multicellular, often sunken and with unicellular or multicellular stalk and multicellular head; secretory hairs (colleters?) present in some species of Ardisia.

Leaves Usually alternate (spiral; sometimes opposite or verticillate), simple, entire, often coriaceous (in Coris ericoid), usually with involute, supervolute or conduplicate (rarely revolute) ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles? Venation usually pinnate (rarely palmate). Stomata anomocytic or anisocytic (rarely helicocytic). Cuticular wax crystalloids as irregular granules (Aegiceras). Lamina gland-dotted or glandular hairy, sometimes with extrafloral nectaries. Epidermis often with mucilage cells. Schizogenous secretory cavities with reddish-brown, red or yellow resin usually present. Mesophyll sometimes with sclerenchymatous idioblasts. Calciumoxalate as druses and solitary crystals present in most tropical woody representatives. Domatia as pockets or hair tufts, or absent. Leaf margin usually entire (sometimes serrate or crenate).

Inflorescence Terminal or axillary, spike-, raceme- or umbel-like, corymbose, fasciculate etc., or flowers solitary. Floral prophylls (bracteoles) absent.

Flowers Usually actinomorphic (in Coris zygomorphic; calyx in Aegiceras asymmetrical). Epicalyx (consisting of ten to 15 spine-shaped acute outgrowths below calyx) present in Coris. Hypogyny. Sepals (three or) four or five (to nine), usually with imbricate (rarely contorted or valvate) aestivation, free or connate (median sepal in Coris abaxial). Petals (three or) four or five (to nine), usually with imbricate or contorted (rarely valvate or conduplicate) aestivation, usually connate into campanulate or urceolate corolla (sometimes only at base; corolla lobes in Cyclamen strongly recurved; in Embelia and Heberdenia free; absent in some species of Lysimachia [’Glaux’]). Petals and stamens developing from common primordia (stamens sometimes initiated as adaxial appendages). Nectariferous hairs often present on petals and carpels; base of ovary in Coris with nectary. Disc?

Androecium Stamens (three or) four or five (to nine), obhaplostemonous, alternisepalous, antepetalous. Filaments free or connate; usually adnate to corolla tube (epipetalous). Anthers dorsifixed or basifixed, sometimes versatile, tetrasporangiate (in Aegiceras transversely septate, locellate), introrse, usually longicidal (dehiscing by longitudinal slits; rarely poricidal, dehiscing by apical pore-like slits), usually free (often connivent into cone-shaped structure around style; rarely connate). Tapetum secretory, with uninucleate cells. Female flowers often with staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually 3(–5)-colporate or 3(–5)-colpate (rarely pantocolporate), shed as monads, bicellular at dispersal. Exine tectate or semitectate, with columellate infratectum, reticulate, scabrate or psilate.

Gynoecium Pistil composed of three to five (or six) connate antepetalous carpels. Ovary superior, unilocular, nectariferous. Style usually single, simple (sometimes absent). Stigma punctate, capitate or truncate (rarely discoid or fimbriate), papillate, Dry or Wet type. Male flowers often with pistillodium?

Ovules Placentation free central to basal. Ovules several (in Coris five or six) to numerous per ovary, usually anatropous (in Cyclamen campylotropous, in Coris hemianatropous), ascending to horizontal, usually bitegmic (in Aegiceras and Cyclamen unitegmic), usually tenuinucellar (in Aegiceras crassinucellar?). Micropyle usually bistomal (in Coris endostomal). Outer integument two or three cell layers thick. Inner integument two or three (to seven) cell layers thick. Integument in Aegiceras four or five cell layers thick, in Cyclamen approx. two cell layers thick. Parietal tissue approx. four cell layers thick (Cyclamen). Archespore multicellular. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria usually absent (in Aegiceras developed from secondary endosperm tissue and invading funicle and integument). Embryogenesis usually caryophyllad (rarely onagrad). Polyembryony sometimes present.

Fruit A drupe, berry or capsule, often dehiscing irregularly (in some species of Lysimachia a pyxidium). Placentae juicy.

Seeds Aril absent. Funicle in Aegiceras not vascularized. Testa multiplicative. Exotesta? Endotestal cells in Cyclamen with crystals and U-shaped wall thickenings. Tegmen at first thick, finally crushed. Endotegmen sometimes with crystals. Perisperm not developed. Endosperm usually copious (often ruminate; absent in Aegiceras), oily and amyloidiferous (in Cyclamen with thick and pitted cell walls). Embryo usually straight (sometimes curved; in Cyclamen diagonal; in Aegiceras very large), without chlorophyll. Cotyledons usually two (in Cyclamen one), short (in Aegiceras connate into tube around plumule). Germination phanerocotylar or cryptocotylar (vivipary present in Aegiceras).

Phytochemistry Flavonols (kaempferol, quercetin, myricetin) and their glycosides, cyanidin, delphinidin, alkaloids, triterpene saponins, and benzoquinones present. Ellagic acid and cyanogenic compounds not found.

Engler in Engler et Prantl, Nat. Pflanzenfam., Nachtr. 1: 214. Oct 1897, nom. cons.

Distribution Southeastern China, northern Indochina, the Malay Peninsula, Sumatra.

Fossils Uncertain. Fossil fruits assigned to Pentaphylax are described from the Maastrichtian of Central Europe, yet their affinity is questioned.

Vegetative anatomy Phellogen? Pericyclic envelope consisting of fibres alternating with lignified parenchyma cells. Vessel elements with scalariform perforation plates; lateral pits opposite to scalariform, simple and/or bordered pits. Imperforate tracheary xylem elements tracheids usually with bordered (sometimes simple) pits, non-septate. Wood rays uniseriate or multiseriate, heterocellular? Axial parenchyma apotracheal diffuse, diffuse-in-aggregates or in short tangential lines. Articulated laticifers abundant. Sieve tube plastids S type? Nodes? Druses absent. Crystals?

Leaves Leaves alternate (spiral or distichous), simple, entire or lobed, with supervolute ptyxis. Stipules and leaf sheath absent. Buds perulate (perulae present). Petiole vascular bundle transection annular. Venation pinnate, often indistinct. Stomata usually paracytic. Cuticular wax crystalloids? Lamina with blackish, caducous and probably glanduliferous apex. Leaf margin serrate or entire.

Inflorescence Up to c. 15 flowers in terminal or axillary racemose inflorescences, with flowers principally solitary in axils of reduced leaves; when floral shoot strongly reduced, then inflorescence becoming fasciculate. Bracts with black, caducous and probably glanduliferous apex. Floral prophylls (bracteoles) two.

Flowers Actinomorphic. Hypogyny. Sepals five, with imbricate quincuncial aestivation, free or more or less connate. Some sepals with black, caducous and probably glanduliferous apex. Petals five, with imbricate quincuncial aestivation, free or more or less connate. Tepals with well developed and distinct canals. Nectary present on ovary wall. Disc absent.

Androecium Stamens five, haplostemonous, alternisepalous, antepetalous. Filaments very wide and relatively long, narrowing distally and incurved at apex, free, tightly pressed against petals and forming tube. Anthers basifixed, non-versatile?, tetrasporangiate, introrse, valvicidal (dehiscing by uplifted subapical valve). Tapetum secretory. Female flowers with staminodia?

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolporate, shed as monads, bicellular at dispersal. Exine tectate, with intermediate infratectum (columellae poorly developed), smooth. Tectum thin. Endexine thick.

Gynoecium Pistil composed of five connate antepetalous carpels. Ovary superior, unilocular. Style single, simple, hollow. Stigmatic areas shortly radiate, type? Male flowers with pistillodium?

Ovules Placentation apical-axile. Ovules two per carpel, campylotropous to hemitropous, pendulous, bitegmic, crassinucellar. Micropyle ?-stomal. Outer integument approx. three? cell layers thick. Inner integument ? cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development? Endosperm haustoria? Embryogenesis?

Fruit A loculicidal capsule, with midrib separating from rest of valves. Endocarp cells transversely elongate.

Seeds Seeds flattened Aril absent. Seed coat winged. Exotestal cells somewhat thickened, with theoid exotestal thickenings, elongate. Mesotestal cells large, thin-walled. Endotesta? Tegmen? Perisperm not developed. Endosperm sparse. Embryo straight?, chlorophyll? Cotyledons longer than radicula. Germination phanerocotylar.

Polemoniales Juss. ex Bercht. et J. Presl, Přir. Rostlin: 248. Jan-Apr 1820 [‘Polemoniae’]; Cobaeaceae D. Don in Edinburgh Philos. J. 10: 111. Jan 1824 [’Cobeaceae’]; Cobaeineae Link, Handbuch 1: 822. 4-11 Jul 1829 [‘Cobaeaceae’]; Polemoniineae Bessey in C. K. Adams, Johnson’s Universal Cyclop. 8: 463. 15 Nov 1895 [‘Polemoniales’]

Distribution Temperate and arctic Eurasia, the Himalayas, North America including arctic areas, Mexico, Central America, northern and western South America to southern Chile and southern Argentina, with their largest diversity in western North America.

Fossils The herbaceous fossil Gilisenium hueberi from the mid-Eocene of Utah has been assigned to Polemoniaceae.

Habit Bisexual, usually perennial, biennial or annual herbs (sometimes shrubs or suffrutices; rarely lianas; in Cantua trees). Often evil-smelling.

Vegetative anatomy Phellogen ab initio usually superficial (sometimes pericyclic). Cambium usually not storied. Endodermis sometimes prominent. Vessel elements usually with simple (rarely also scalariform) perforation plates; lateral pits alternate, bordered pits. Imperforate tracheary xylem elements tracheids, fibre tracheids or libriform fibres with simple or bordered pits, septate or non-septate. Wood rays uniseriate or multiseriate, heterocellular, or absent. Axial parenchyma paratracheal scanty vasicentric. Wood elements often storied. Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace. Phloem in Cobaea with secretory cavities containing yellow secretion. Crystals?

Trichomes Hairs multicellular, usually uniseriate (in, e.g., Langloisia branched), sometimes arachnoid, bristle-shaped or dendritic (sometimes calcified/silicified); glandular hairs with unicellular or multicellular head often present.

Leaves Usually alternate (spiral; in Linanthus and Phlox opposite; in Gymnosteris verticillate), simple or pinnately or palmately compound, entire or lobed (in Cobaea with terminal leaflet modified into tendril; in Acanthogilia dimorphic, with young leaves pinnately compound and older leaves simple and entire), with conduplicate ptyxis. Stipules usually absent (in Cobaea intrapetiolar, foliaceous); leaf sheath absent. Colleter-like glandular hairs often present on adaxial side of petiole base. Petiole vascular bundles? Venation usually pinnate (sometimes palmate). Stomata usually anomocytic (sometimes almost paracytic). Cuticular wax crystalloids? Epidermis with or without mucilage cells. Leaf margin lobed, serrate or entire. Multicellular bristles present in Langloisia and some species of Loeselia.

Inflorescence Usually terminal (in Acanthogilia axillary), umbel-like, capitate, corymbose, panicle, thyrsopaniculate or botryoid (flowers rarely solitary). Inflorescences sometimes pseudanthia with involucral bracts. Floral prophylls (bracteoles) absent.

Flowers Usually actinomorphic (sometimes zygomorphic). Hypogyny. Sepals (four or) five (or six), with imbricate, valvate or open aestivation, persistent, connate (rarely almost free). Petals (four or) five (or six), usually with convolute or contortuplicate (rarely imbricate) aestivation, connate into campanulate, infundibuliform or discoid corolla; median petal adaxial or abaxial; corolla sometimes bilabiate (e.g. 2:3, 3:2 or 5:0). Nectariferous disc intrastaminal, annular, entire or lobed (rarely absent).

Androecium Stamens (three to) five (or six), usually as many as sepals, haplostemonous, antesepalous, alternipetalous, often unequal in length. Filaments free from each other, usually adnate to corolla tube (epipetalous; sometimes free from petals). Anthers usually ventrifixed (rarely dorsifixed), in Cobaea versatile, tetrasporangiate, introrse?, longicidal (dehiscing by longitudinal slits). Tapetum usually secretory (in Cobaea amoeboid-periplasmodial?). Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually (tetra- to) hexa- to c. 50-pantoporate (rarely zonotreme, bizonotreme, anomotreme/porate or [stephano]colp[or]ate; in Acanthogilia zonotreme), shed as monads, bicellular at dispersal. Exine tectate to semitectate, with columellate infratectum, reticulate, striate or verrucate.

Gynoecium Pistil composed of (two or) three (or four) connate carpels, median carpel adaxial. Ovary superior, (bilocular or) trilocular (or quadrilocular). Style single, filiform, usually trilobate. Stigmas decurrent along entire lobes, papillate, Dry type. Pistillodium absent.

Ovules Placentation axile. Ovules one to numerous per carpel, anatropous to hemitropous, apotropous, unitegmic, usually tenuinucellar (in Cobaea pseudocrassinucellar). Integument three to 20 cell layers thick. Nucellar cap present at least in Cobaeoideae. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear (endosperm finally cellular). Endosperm haustoria absent. Embryogenesis chenopodiad.

Fruit Usually a loculicidal (in Acanthogilia and Cobaea septicidal) capsule (rarely a nut).

Seeds Aril?Seed coat sometimes winged, usually mucilaginous when wet. Exotesta of various thickness, with thickened radial cell walls. Epidermal cells often cylindrical with distinct spiral wall thickenings and mucilage. Endotesta reduced to pigmentary layer. Perisperm not developed. Endosperm usually copious (sometimes sparse), oily (absent in Cobaea). Embryo usually straight (sometimes curved), well differentiated (Cobaea), with or without chlorophyll. Cotyledons two. Germination usually phanerocotylar (in Cobaea cryptocotylar).

Cytology n = 6–12, with large chromosomes (Polemonioideae); n = (9, Acanthogilia) 15, 26, 27, with small chromosomes (Cobaeoideae) – Polyploidy and aneuploidy occurring.

Phytochemistry Flavonols (kaempferol, quercetin), 6-metoxyflavonols, flavone-C-glycosides etc., cyanidin, oleanolic acid derivatives, triterpene saponins, and cucurbitacins present. Iridoids, ellagic acid, alkaloids, and cyanogenic compounds not found. Carbohydrates stored as oligosaccharides with kestose and isokestose bonds (inulin).

Systematics Polemoniaceae are sister to Fouquieriaceae.

Cobaeoideae Arn., Botany: 121. 9 Mar 1832 [‘Cobaeeae’]

3/31–33. Cantua (12; the Andes in Ecuador, Peru and Bolivia), Cobaea (18–20; Mexico, Central America, tropical South America), Bonplandia (1; B. geminiflora; Mexico, Guatemala). – Baja California in northwestern Mexico to tropical South America. Twining herbs, lianas, trees. Stipules in Cobaea intrapetiolar, foliaceous. Flowers large, in Cobaea somewhat zygomorphic. Sepals largely connate (sometimes only at base), with green midrib. Petal veins united at base of lobe (sometimes also upper lobe). Staminal traces present in two whorls. Filaments often superficially adnate to corolla. Pollen grains sometimes 100–220 μm, with verrucate exine. Nucellar cap present. Fruit a septicidal and/or loculicidal capsule. Seed coat narrowly or broadly winged. Mesotestal cell walls thinly lignified. n = 15, 26, 27. – Cobaeoideae may be sister-group to [Acanthogilioideae+Polemonioideae] (Johnson & al. 2008).

Acanthogilioideae J. M. Porter et L. A. Johnson in Aliso 19: 60. 28 Jul 2000

1/1. Acanthogilia (1; A. gloriosa; Baja California in northwestern Mexico). – Shrub. Leaves very dimorphic, deciduous on short shoots, modified into branched spines on long shoots. Sepals with green midrib. Pollen grains stephanocolporate. Exine corsely verrucate. Seeds few. n = 9. – Acanthogilia is sister to either the clade [Cobaeoideae+Polemonioideae], Cobaeoideae or Polemonioideae. Johnson & al. (2008) recovered Acanthogilia as sister to Polemonioideae.

Polemonioideae Arn., Botany: 121. 9 Mar 1832 [‘Polemonieae’]

19/345–350. Western North America, northern temperate regions, southern South America. Usually herbs (sometimes suffrutices). Flowers sometimes zygomorphic. Petal veins usually free or united high above base. Filaments usually adnate to corolla tube (free from petals in, e.g., Collomia and Loeselia. Hypostase present. Seed coat usually not winged (in Loeselia narrowly winged). Testa sometimes multiplicative. n = (6) 7 (8) 9 (or more). – A plausible topology is [[Polemonieae+Phlocideae]+[Gilieae+Loeselieae]] with high posterior probability but low bootstrap support (Johnson & al. 2008).

[Polemonieae+Phlogieae]

Polemonieae Dumort., Anal. Fam. Plant.: 25. 1829 [‘Polemoneae’].

1/c 30. Polemonium (c 30; temperate regions on the Northern Hemisphere south to Mexico, Chile). – Usually perennial (P. micranthum annual) herbs. Leaves alternate (spiral), pinnate. Calyx accrescent. Seeds dark, shiny. – Polemonium has sometimes been recovered as sister to the remaining Polemonioideae, to Phlocideae, or to [Gilieae+Phlocideae].

Phlocideae J. M. Porter et L. A. Johnson in Aliso 17: 84. 8 Jun 1998 (Phlocideae Dumort., Anal. Fam. Plant.: 25. 1829).

3/120–125. Linanthus (c 55; southwestern Canada, western United States, Mexico), Gymnosteris (2; G. nudicaulis, G. parvula; western United States), Phlox (65–70; northeastern Asia, Canada, United States, northern Mexico, one species in Siberia). – Northeastern Asia, North America to northern Mexico. Leaves usually opposite. Corolla usually hypocraterimorphous. – Linanthus is sister to the remainder.

[Loeselieae+Gilieae]

Loeselieae J. M. Porter et L. A. Johnson in Aliso 17: 84. 8 Jun 1998.

9/c 97. Aliciella (21; western United States, northwestern Mexico), Giliastrum (10; western United States, Mexico, Central America), Loeselia (14; California), Dayia (5; D. glutinosa, D. grantii, D. havardii, D. scabra, D. sonorae; southern United States, northern and northwestern Mexico, northern Chile), Bryantiella (1; B. palmeri; southern California; in Ipomopsis?), Ipomopsis (c 30; southwestern Canada, United States, Mexico, Chile, Argentina; non-monophyletic?), Microgilia (1; M. minutiflora; western United States), Eriastrum (14; southwestern United States), Langloisia (1; L. setosissima; southwestern United States, northern Mexico). – Style usually persistent in fruit. – Aliciella is sister-group to the remaining Loeselieae.

Gilieae (Rchb.) V. E. Grant, Nat. Hist. Phlox Fam.: 120. 1959.

6/c 100. Saltugilia (4; S. australis, S. caruifolia, S. latimeri, S. splendens; California, Baja California); Lathrocasis (1; L. tenerrima; western United States, northwestern Mexico), ’Gilia’ (c 40; southwestern Canada, western United States, Mexico to northern Chile; non-monophyletic), Allophyllum (5; A. divaricatum, A. gilioides, A. glutinosum, A. integrifolium, A. nemophilophyllum; western United States), Collomia (15; southwestern Canada, western United States, Mexico, Bolivia to temperate Chile and Argentina), Navarretia (35; southwestern Canada, western United States, northern Mexico, Chile, Argentina). – Saltugilia is sister to the rest.

Cladogram of Polemoniaceae based on DNA sequence data (Johnson & al. 2008). Acanthogilia is sometimes recovered as sister to Cobaeoideae and/or Polemonioideae.

Primulales Juss. ex Bercht. et J. Presl, Přir. Rostlin: 241. Jan-Apr 1820 [‘Primulaceae’]; Primulineae Burnett, Outlines Bot.: 901, 1101. Feb 1835 [‘Primulosae’]; Primulopsida Brongn., Enum. Plant. Mus. Paris: xxi, 69. 12 Aug 1843 [’Primulineae’]; Hottoniaceae Döll, Fl. Baden 2: 641. med 1858 [’Hottonieae’]

Distribution Temperate and arctic-alpine regions on the Northern Hemisphere, tropical mountains in East Africa, the Arabian Peninsula and Malesia, southernmost South America.

Fossils Fossil long-stalked ebracteate hypogynous to almost hypogynous pentamerous flowers of primuloid origin were described from the Campanian to the Maastrichtian of Portugal. The ovary is unilocular and the placentation is free central, with numerous anatropous ovules. Most of the outer tissues are beset with schizogenous secretory cavities.

Habit Bisexual, perennial or annual herbs (sometimes cushion-shaped and woody at base), usually with a basal leaf rosette. Some species are xerophytes. Hottonia is aquatic.

Vegetative anatomy Phellogen? Primary vascular tissue a cylinder without separate vascular bundles, or a cylinder of bundles or scattered bundles. Secondary lateral growth anomalous or absent. Endodermis usually prominent. Vessel elements with simple perforation plates; lateral pits alternate. Imperforate tracheary xylem elements libriform fibres with simple or bordered pits. Wood rays multiseriate, heterocellular? Axial parenchyma? Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace. Secretory cells and schizogenous resinous canals present. Crystals?

Trichomes Hairs multicellular, uniseriate, articulated, often long, stellate hairs present in some species of Androsace and Primula; glandular hairs short; immersed glandular hairs absent.

Leaves Usually alternate (sometimes opposite or verticillate), usually simple (rarely pinnately compound), usually entire (rarely pinnately lobed), with involute or conduplicate (sometimes revolute) ptyxis. Stipules or leaf sheath absent. Petiole vascular bundles? Venation pinnate or palmate; marginal veins ending with a hydathode. Stomata usually anomocytic. Cuticular wax crystalloids as threads, chemically dominated by flavonoids. Lamina in Omphalogramma, Bryocarpum and some species of Primula with black flattened glandular dots. Sclereids present in some species. Schizogenous or lysigenous secretory cavities? Leaf margin serrate, crenate or entire.

Inflorescence Terminal or axillary, verticels, a single umbel, raceme, spike, or head, or cymose and usually long-stalked (scapose), or flowers solitary. Floral prophylls (bracteoles) absent.

Flowers Usually actinomorphic (rarely somewhat zygomorphic). Hypogyny. Sepals five (to eight), usually with imbricate aestivation, usually persistent, often connate. Petals five (to eight), usually with imbricate aestivation, connate into campanulate or discoid (hypocraterimorphous) corolla (corolla lobes in Soldanella fimbriate, in some species of Primula [’Dodecatheon’] strongly recurved). Petals and stamens developing from common primordia. Nectaries present on ovary. Disc absent. Diheterostyly frequent.

Androecium Stamens five (to eight), haplostemonous, antepetalous, alternisepalous. Filaments free, adnate to corolla tube (epipetalous). Anthers sometimes connivent and strongly protruding, dorsifixed or basifixed, non-versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, with uninucleate cells. Staminodia usually absent (in Soldanella extrastaminal, antesepalous, scale-like).

Pollen grains Microsporogenesis simultaneous. Pollen grains (2–)3(–10)-colporate, -colporoidate or -colpate (in many species of Primula syncolpate or stephanocolpate), shed as monads, bicellular at dispersal. Exine tectate, with columellate infratectum, perforate, often granulate.

Gynoecium Pistil composed of five connate antepetalous (in Primula antesepalous?) carpels. Ovary superior, unilocular (rarely with rudimentary septa at base), nectariferous. Style single, simple, with stylar canal. Stigma truncate to capitate, papillate or non-papillate, Dry type. Pistillodium absent.

Ovules Placentation free central. Ovules five to more than 100 per ovary, anatropous or hemitropous, ascending, usually bitegmic (in Androsace [‘Douglasia’] unitegmic), tenuinucellar. Micropyle bistomal. Outer integument approx. two cell layers thick. Inner integument three or four cell layers thick. Integument in ‘Douglasia’ six to ten cell layers thick. Archespore multicellular. Megagametophyte monosporous, Polygonum type Endosperm development ab initio nuclear. Endosperm haustoria absent. Embryogenesis caryophyllad. Polyembryony present in some species.

Fruit A denticidal or irregularly dehiscing capsule (in one? species of Androsace [’Pomatosace’] a pyxidium).

Seeds Aril absent. Exotesta persistent; exotestal cell walls often thick. Endotesta in Primula with thick inner cell walls. Exotegmen? Endotegmen often with rhomboidal crystals. Perisperm not developed. Endosperm usually copious, usually with thick and pitted cell walls, oily and amyloidiferous (absent in Hottonia and Soldanella). Embryo short, straight, without chlorophyll. Cotyledons two, short. Germination phanerocotylar.

DNA Plastid gene infA lost/defunct (Primula). Two deletions in plastid gene ndhF.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), cyanidin, oleanolic acid derivatives, ellagic acid (at least in Hottonia), proanthocyanidins (prodelphinidins), alkaloids, triterpene saponins, benzoquinones (rapanone), and cucurbitacins (absent from Soldanella) present. Cyanogenic compounds not found.

Roridulales Bercht. et J. Presl, Přir. Rostlin: 217. Jan-Apr 1820 [‘Roridulae’]

Habit Bisexual, evergreen shrubs. Semicarnivorous (producing resins without digesting enzymes); living in mutualistic relationship with two species of carnivorous hemipterans, Pameridea marlothii (on Roridula dentata) and P. roridulae (on both species of Roridula), which feed on insects trapped by the sticky resins from the glandular hairs. The urea excreted by the bugs is absorbed by the Roridula plant through the leaf surface. The Pameridea species have not been found in nature outside Roridula.

Vegetative anatomy Mycorrhiza absent? Phellogen? Stem endodermis continuous and starchy; inside this a sclerenchymatous ring. Primary medullary rays narrow. Pericyclic fibres absent. Vessel elements with scalariform or reticulate perforation plates; lateral pits usually alternate, scalariform or opposite, bordered pits. Imperforate tracheary xylem elements tracheids with bordered pits. Wood rays usually uniseriate (rarely biseriate or multiseriate), homocellular. Axial parenchyma apotracheal diffuse, or paratracheal scanty. Sieve tube plastids Ss type. Nodes? Wood ray cells sometimes with gum-like deposits. Calciumoxalate druses sometimes present.

Trichomes Foliar hairs unicellular bristle-like and multicellular glandular hairs; glandular hairs stalked, capitate, resin-producing; gland stalk uniseriate to quinqueseriate; head sexacellular to multicellular, in larger glandular hairs with central column bearing resin-producing cells.

Leaves Alternate (spiral), simple, entire to narrowly pinnately lobed (lamina linear, narrowing towards apex), with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles? Venation brochidodromous. Stomata anomocytic. Cuticular wax crystalloids? Epidermis with calciumoxalate druses. Leaf margin entire or narrowly lobate. Resin-producing glandular hairs numerous along margin and along midvein on abaxial side, also elsewhere.

Inflorescence Terminal, botryoid or raceme-like cymose, or flowers solitary axillary.

Flowers Actinomorphic. Hypogyny. Sepals five, with quincuncial imbricate aestivation, persistent, free (connate at base?). Petals five, with quincuncial imbricate aestivation, free (slightly connate at base?). Nectaries present on staminal bases. Disc absent.

Androecium Stamens five, haplostemonous, antesepalous, alternipetalous. Filaments filiform, thicker at base, free, adnate to petal base, nectariferous. Anthers subbasifixed, in bud inflexed, early inverted (as mature resupinated), versatile?, tetrasporangiate, seemingly extrorse, poricidal (dehiscing by apical pores or short slits); connective swollen at anther base. Fibrous endothecium absent. Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolpor(oid)ate (rarely tetracolporate), shed as monads, bicellular at dispersal. Exine tectate, with granular infratectum, perforate, spinulate (Roridula dentata) or insular with angular to rounded segments (Roridula gorgonias).

Gynoecium Pistil composed of three connate carpels. Ovary superior, trilocular. Style single, simple. Stigma capitate or reversed conical, papillate, type? Pistillodium absent.

Ovules Placentation axile to apical. Ovules one to four per carpel, anatropous, pendulous, unitegmic, tenuinucellar. Integument ? cell layers thick. Hypostase? Megagametophyte monosporous, Polygonum type. Endosperm development cellular. Endosperm haustorium micropylar. Embryogenesis solanad.

Fruit A loculicidal (and septicidal?) capsule, with central columella persistent after dehiscence.

Seeds Aril? Exotesta mucilaginous when moistened; outer epidermal cells of exotesta with thickened inner tangential and radial walls. Endotesta? Perisperm not developed. Endosperm copious, fleshy. Embryo small, straight, well differentiated, chlorophyll? Cotyledons two. Germination phanerocotylar.

Phytochemistry Insufficiently known. Route I iridoids (also secoiridoids?) present. Tannins present in endosperm and integument. Ellagic acid? Proanthocyanidins, cyanogenic compounds and naphthoquinones not found.

Rafinesque in Ann. Gén. Sci. Phys. Bruxelles 5: 349. Jul-Sep 1820 [’Samolia’]

Distribution Europe, Macaronesia, Africa, southwestern Asia to southern China, East Asia to Japan, southern Australia, Tasmania, New Caledonia, New Zealand, eastern and southern North America, southern and central South America.

Habit Bisexual, usually perennial or annual herbs (sometimes woody at base).

Vegetative anatomy Phellogen? Vessel elements with simple? perforation plates; lateral pits alternate? Imperforate tracheary xylem elements libriform fibres? Wood rays absent? Axial parenchyma? Sieve tube plastids S type? Nodes 1:1?, unilacunar with one leaf trace? System of secretory canals present especially in inflorescence. Crystals?

Trichomes Hairs usually absent; sessile or stalked glandular hairs with one or few stalk cells and unicellular or multicellular head sometimes present.

Leaves Alternate (spiral), simple, entire (rarely scale-like), with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate; bundles probably diverging very soon after leaf trace departs from central stele. Venation pinnate. Stomata anomocytic. Cuticular wax crystalloids? Leaf margin entire.

Inflorescence Terminal, simple or compound raceme or corymbose. Floral prophylls (bracteoles) absent.

Flowers Actinomorphic. Half epigyny (perigyny). Sepals five, with open aestivation, persistent, connate, adnate to ovary. Petals five, with imbricate aestivation, connate into campanulate or urceolate corolla. Petals and stamens developing from common primordia; stamen primordia initially larger. Nectaries present on ovary. Disc absent.

Androecium Stamens five, haplostemonous, antepetalous, alternisepalous. Filaments free, adnate to corolla tube (epipetalous). Anthers basifixed, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits); connective sometimes prolonged. Tapetum secretory? Staminodia five, extrastaminal, antesepalous, alternipetalous, triangular or linear, epipetalous, or absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolporoidate, shed as monads, bicellular at dispersal. Exine tectate, with columellate? infratectum, finely reticulate.

Gynoecium Pistil composed of five connate antepetalous carpels. Ovary semi-inferior, unilocular, nectariferous. Style single, simple, impressed. Stigma truncate to capitate, type? Pistillodium absent.

Ovules Placentation free central to basal. Ovules numerous per ovary, hemitropous, bitegmic, tenuinucellar. Micropyle bistomal? Outer integument approx. two cell layers thick. Inner integument approx. two cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria absent? Embryogenesis caryophyllad?

Fruit A many-seeded, loculicidal-denticidal, quinquedentate capsule with five antesepalous valves.

Seeds Aril absent? Seed coat indistinct. Exotestal cells tanniniferous. Endotesta? Exotegmen? Endotegmen tanniniferous, with rhomboidal crystals. Perisperm not developed. Endosperm copious, with thin to slightly thickened cell walls, amyloidiferous. Embryo small, straight, chlorophyll? Cotyledons two, narrow. Germination phanerocotylar?

Phytochemistry Insufficiently known. Flavonols (kaempferol, quercetin, myricetin)? and other polyphenols present. Cyanidin? Delphinidin? Glycosides? Triterpene saponins? Ellagic acid, alkaloids and cyanogenic compounds not found?

Achradaceae Vest, Anleit. Stud. Bot.: 273, 299. 1818 [’Achratoideae’]; Sapotales Juss. ex Bercht. et J. Presl, Přir. Rostlin: 250. Jan-Apr 1820 [‘Sapoteae’]; Bumeliaceae Barnhart in Bull. Torrey Bot. Club 22: 21. 15 Jan 1895; Sapotineae Engl., Syllabus, ed. 2: 169. Mai 1898; Boerlagellaceae H. J. Lam in Bull. Jard. Bot. Buitenzorg, sér. 3, 7: 250. Feb 1925; Sarcospermataceae H. J. Lam in Bull. Jard. Bot. Buitenzorg, sér. 3, 7: 248. Feb 1925 [’Sarcospermaceae’], nom. cons.

Distribution Mainly pantropical; some species in subtropical regions (southeastern United States, central South America, southeastern Africa, South Asia).

Fossils Pollen resembling those in Sapotaceae have been reported from Eocene and younger layers. Late Oligocene branches and fruits of putative sapotacean origin are known from Germany.

Habit Usually bisexual (sometimes monoecious or dioecious, rarely gynomonoecious), evergreen trees or shrubs (rarely lianas).

Vegetative anatomy Phellogen ab initio superficial. Medullary vascular bundles sometimes present. Vessel elements usually with simple (rarely scalariform) perforation plates; lateral pits alternate, usually simple pits. Imperforate tracheary xylem elements fibre tracheids or libriform fibres usually with simple (sometimes bordered) pits, usually non-septate (in Sarcosperma also septate; vasicentric tracheids abundant). Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal reticulate, scalariform or banded. Tyloses often frequent (sometimes sclerotic). Sieve tube plastids S type. Nodes usually 3:3, trilacunar with three leaf traces (rarely 1:1, unilacunar with one trace). Laticifers articulated, usually with white (rarely yellow or bluish-green) latex. Wood elements often silicified and/or with silica bodies. Calciumoxalate crystals as druses, styloids, acicular and prismatic crystals and/or crystal sand sometimes present.

Trichomes Hairs usually unicellular (in Delpydora multicellular), uniseriate or T-shaped, often malpighiaceous, with branches of unequal lengths, often brownish.

Leaves Usually alternate (spiral or distichous; sometimes opposite or verticillate), simple, entire, coriaceous, with conduplicate ptyxis. Stipules cauline (intrapetiolar) or absent; leaf sheath absent. Petiole often swollen at base (in some species of Sarcosperma and Pradosia with two stipule-like appendages, stipulules, at apex). Petiole vascular bundle transection arcuate, D-shaped or annular; petiole sometimes with wing bundles. Venation pinnate, usually brochidodromous (sometimes eucamptodromous or craspedodromous); secondary veins often parallel and densely spaced. Stomata usually anomocytic (sometimes paracytic). Cuticular wax crystalloids? Epidermis with or without mucilage cells. Mesophyll usually with sclerenchymatous idioblasts; mesophyll fibres sometimes with spiral thickenings. Special laticifers and cells secreting gutta-percha. Sclereids present. Domatia as pits. Leaf margin usually entire (sometimes serrate, rarely serrate-dentate). Two-branched hairs frequent.

Inflorescence Axillary, usually fasciculate, raceme-like or panicle (sometimes solitary axillary).

Flowers Actinomorphic. Pedicel not articulated. Hypogyny. Sepals four to six, in one whorl, usually with imbricate quincuncial aestivation, free or connate in lower part; or six to eleven, spiral, with imbricate aestivation, free; or 2+2, 3+3 or 4+4, outer sepals with valvate, inner sepals with imbricate aestivation, free or connate at base. Petals three to six, 4+4, 5+5 or up to 18, entire, lobed or divided above, or divided down to base into three segments with mid-segment entire and two lateral (dorsal) segments entire, lobed or connate. Nectariferous disc annular or cushion-shaped, or absent.

Androecium Stamens four to c. 35(–43), as many as petals, alternisepalous, antepetalous; or (when more numerous than petals) some stamens antepetalous and others alternipetalous; or many stamens in antepetalous fascicles; or numerous stamens (rarely up to six times as many as petals) in two or three whorls. Filaments usually free (rarely connate into a tube), usually adnate to corolla tube or corolla lobes. Anthers basifixed or dorsifixed, sometimes versatile?, tetrasporangiate (in Magodendron locellate, with septa), introrse to extrorse (in Sarcosperma sometimes latrorse), longicidal (dehiscing by longitudinal slits). Tapetum secretory, with binucleate to quadrinucleate cells. Staminodia up to eight (to twelve), extrastaminal, in one whorl, antesepalous, alternating with stamens, often adnate to corolla tube, often lobed or divided (sometimes petaloid), or absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains (2–)3–4(–6)-colporate, shed as monads, bicellular or tricellular at dispersal. Exine tectate, with columellate or granular infratectum, punctate, perforate, microreticulate, reticulate, psilate, scabrate, verruculate, striate or rugulate (rarely spinulate).

Gynoecium Pistil composed of (one to) four to 14 (to 30) connate antesepalous carpels. Ovary superior (unilocular to) quadrilocular to 14-locular (to 30-locular), with hairs inside. Style single, intire. Stigma punctate or somewhat lobate, papillate, Dry type. Pistillodium?

Ovules Placentation usually axile or axile-basal (when ovary unilocular, then basal-parietal). Ovules one (to five) per carpel, anatropous to hemianatropous, ascending, apotropous, unitegmic, tenuinucellar. Integument ? cell layers thick. Hypostase absent (not organized). Megagametophyte monosporous, Polygonum type. Synergids at least sometimes with a filiform apparatus. Antipodal cells ephemeral. Endosperm development ab initio nuclear. Endosperm haustorium absent. Embryogenesis?

Fruit Usually a berry (rarely a drupe or a loculicidal capsule) with persistent calyx.

Seeds Aril absent. Testa multiplicative. Exotestal cells isodiametric and strongly lignified and sclerotized (tanniniferous). Endotesta finally crushed. Perisperm not developed. Endosperm copious to sparse, oily, with or without amyloid?, or absent. Embryo straight, well differentiated, without chlorophyll. Cotyledons two, foliaceous. Germination phanerocotylar or cryptocotylar.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), cyanidin, oleanolic acid derivatives, C-30-oxidized triterpenes, pentacyclic triterpenes, latex of trans- and/or cis-polyisoprenes, dammaranes, gallic acid, condensed and hydrolyzable tannins, proanthocyanidins (prodelphinidins), pyrrolizidine alkaloids as aliphatic monocarboxylic esters and esters of arylic and aralkylic acids, saponins, tyrosine- or phenylalanine-derived cyanogenic compounds naphthoquinones, dihydrosterculic acid, myo-inisitol, and resins present. Ellagic acids not found.

Use Fruits (Manilkara, Chrysophyllum), latex (gutta-percha, balata, chicle, tewing gums), seed oil (Butyrospermum), timber.

Systematics Sapotaceae may be sister to the clade [Ebenaceae+[Maesa+[Samolus+ Theophrastaceae]+[Myrsinaceae+Primulaceae]]].

Sapotaceae and Ebenaceae have similar malpighiaceous hairs. Sapotaceae and Lecythidaceae have seeds with lignified multiplicative testal cells.

A plausible topology is [Sarcosperma+[Eberhardtia+[Xantolis+[‘Sapotoideae’+’Chrysophylloideae’]]]].

Bayesian majority rule consensus tree of Sapotaceae based on DNA sequence data and morphology (Smedmark & al. 2006).

Sarcospermatoideae (H. J. Lam) Swenson et Anderb. in Cladistics 21: 120. Apr 2005

1/11. Sarcosperma (11; eastern Himalayas, southern China, Southeast Asia, Malesia). – Laticifers and latex sometimes absent? Leaves opposite. Stipules cauline. Some species with paired stipulules on petiole apex. Inflorescence axis well developed (in reality consisting of a reduced branch). Staminodia scale-like, short and relatively wide. Pistil composed of one or two connate carpels. Style stout. Hilar scar distinct. Endosperm absent. – Sarcosperma is sister-group to the remaining Sapotaceae.

[Eberhardtia+[Xantolis+[‘Sapotoideae’+’Chrysophylloideae’]]]

Stipules cauline or absent. Stamens as many as corolla lobes, antepetalous, or twice (to six times) as many as corolla lobes. Staminodia present, often petaloid, antesepalous (sometimes absent). Pistil composed of one to 14 (to 30) connate antesepalous carpels. Seed in Omphalocarpum with amyloid (xyloglucans).

Eberhardtia

1/3. Eberhardtia (3; E. aurata, E. krempfii, E. tonkinensis; southern China, Laos, Vietnam).

[Chrysophylloideae+[Neohemsleya+[[Xantolis+Spiniluma oxyacantha]+Sapotoideae]]]

Chrysophylloideae Leurss., Handb. Syst. Bot. 2: 946. Jul 1882 [‘Chrysophylleae’]

36/660–680. Vanroyena (1; V. castanosperma; northeastern Queensland), Sersalisia (2; S. sericea, S. sessiliflora; tropical Australia), Pichonia (10; the Moluccas, New Guinea, the Bismarck Archipelago, Solomon Islands, New Caledonia), Magodendron (2; M. mennyae, M. venefici; Papua New Guinea), Niemeyera (3; N. chartacea, N. prunifera, N. whitei; Queensland, New South Wales), Pycnandra (c 60; New Caledonia),‘Planchonella’ (c 60; East Malesia to islands in western Pacific; polyphyletic), Amorphospermum (1; A. antilogum; northeastern and eastern Queensland, northeastern New South Wales), Micropholis (c 40?; Central America, tropical South America), Ragala (5; R. balata, R. guianensis, R. sanguinolenta, R. spuria, R. ulei; northeastern South America), Ecclinusa (11; Trinidad, tropical South America), Elaeoluma (4; E. crispa, E. glabrescens, E. nuda, E. schomburgkiana; Panamá to central Brazil), Nemaluma (2; N. anomala, N. engleri; northeastern South America), Pleioluma (>30; Malesia to tropical Australia, New Caledonia), Prieurella (7; tropical South America), Diploon (1; D. cuspidatum; southeastern Brazil), Martiusella (2?; M. gonocarpa, M. imperialis; eastern Brazil), Cornuella (1; C. venezuelanensis; northern South America), Achrouteria (2; A. durifructa, A. pomifera; northeasternmost South America?; in Chloroluma?), Chloroluma (1; C. gonocarpum; tropical South America; incl. Achrouteria?), Chrysophyllum (15–20?; Africa, Madagascar, India, Southeast Asia, Malesia to tropical Australia, tropical America; incl. Villocuspis?), Villocuspis (6?; tropical South America; in Chrysophyllum?), Donella (17; tropical Africa, Madagascar), Gambeya (2–5; G. africana, G. korupensis, G. lacourtiana, G. madagascariensis, G. subnuda; Central Africa), Lucuma (20–25; tropical South America), Sarcaulus (5; S. brasiliensis, S. inflexus, S. oblatus, S. vestitus, S. wurdackii; tropical South America; in Pouteria?), Chromolucuma (8; southern Central America, tropical South America; in Pouteria?), Pradosia (23; tropical South America; in Pouteria?), Pouteria (c 300?; tropical Africa, tropical Asia to the Pacific, tropical America). – Unplaced Chrysophylloideae Aubregrinia (1; A. taiensis; Ivory Coast, Ghana), Breviea (1; B. sericea; Ivory Coast, Ghana, Congo?), Delpydora (2; D. gracilis, D. macrophylla; Gabon), Diploon (1; D. cuspidatum; southeastern Brazil), Englerophytum (10–15; tropical Africa), Omphalocarpum (27; tropical West and Central Africa), Sarcaulus (5; tropical South America), Synsepalum (c 20?; tropical Africa), Tridesmostemon (4; T. bequaertii, T. congoense, T. mortehani, T. omphalocarpoides; Central Africa).

Simplified phylogeny (maximum clade credibility tree) of Chrysophylloideae based on nuclear DNA sequence data (Swenson & al. 2013).

[Neohemsleya+[[Xantolis+Spiniluma oxyacantha]+Sapotoideae]]

Neohemsleya

1/1. Neohemsleya (1; N. usambarensis; Tanzania). – Neohemsleya usambarensis was sister to Sapotoideae in Gautier & al. (2013).

[[Xantolis+Spiniluma oxyacantha]+Sapotoideae]

[Xantolis+Spiniluma oxyacantha]

2?/c 16. Xantolis (c 15; South Asia, southern China, Indochina, the Philippines), Spiniluma (1; S. oxyacantha; Ethiopia, Erithrea, southern Arabian Peninsula). – The sister-group relationship between Xantolis and Spiniluma is uncertain. The only species of Spiniluma investigated by Gautier & al. (2013) was S. oxyacantha (S. discolor is the second species in Spiniluma). They found this clade to be sister-group to Sapotoideae.

Sapotoideae Eaton, Bot. Dict., ed. 4: 35. Apr-Mai 1836

c 23?/515–520. Sideroxyleae Small, Man. S.E. Fl.: 1031. 30 Nov 1933. Sideroxylon (75–80; tropical and subtropical regions on both hemispheres). – Lecomtedoxa (6; L. biraudii, L. heitzana, L. klaineana, L. nogo, L. plumosa, L. saint-aubinii; Gabon), Neolemonniera (3; N. batesii, N. clitandrifolia, N. ogouensis; tropical West Africa). The clade [Lecomtedoxa+Neolemonniera] was sister-group to the remaining Sapotoideae in Gautier & al. (2013). – Tseboneae L. Gaut. Et Naciri in Taxon 62: 979. 22 Oct 2013. Tsebona (1; T. macrantha; Madagascar), Bemangidia (1; B. lowryi; southeastern Madagascar), Capurodendron (23; Madagascar). – Isonandreae T. D. Penn., Gen. Sapotac.: 147. 1991. Burckella (14; East Malesia to Tonga), ’Madhuca’ (c 100; India, Sri Lanka, Southeast Asia, Malesia to New Guinea, tropical Australia; polyphyletic), Diploknema (7; D. butyracea, D. butyraceoides, D. oligomera, D. ramiflora, D. sebifera, D. siamensis, D. yunnanensis; India to Yunnan, Southeast Asia and Borneo), Payena (c 20; West Malesia, Mindanao, with their highest diversity on Borneo; non-monophyletic?), Isonandra (c 10; southern India, Sri Lanka), Palaquium (c 110; India, Sri Lanka, southern China, Southeast Asia, Taiwan, Malesia to Samoa), Northia (1; N. seychellana; the Seychelles). – Sapoteae Reichenb., Handb. Nat. Pflanz.-Syst.: 214. 1–7 Oct 1837. Inhambanella (2; I. guereensis, I. henriquezii; western and southeastern tropical Africa), Vitellariopsis (5; V. cuneata, V. dispar, V. ferruginea, V. kirkii, V. marginata; East Africa), Vitellaria (1; V. paradoxa; tropical West and Central Africa; in Vitellariopsis?), Tieghemella (2; T. africana, T. heckelii; tropical West and Central Africa), Mimusops (c 40; tropical Africa, Madagascar, the Mascarene Islands, the Seychelles, one species, M. elengi, to tropical Asia), Autranella (1; A. congolensis; tropical West and Central Africa), Labramia (9; Madagascar), Labourdonnaisia (7; L. calophylloides, L. glauca, L. lecomtei, L. madagascariensis, L. revoluta, L. richardiana, L. thouarsii; Madagascar, Mauritius, Réunion), Faucherea (11; Madagascar), Manilkara (c 65; tropical regions on both hemispheres).

Bayesian majority rule consensus tree of Sapotoideae based on DNA sequence data (Gautier & al. 2013).

Unplaced Sapotaceae

Baillonella (1; B. toxisperma; Central Africa to Angola), Gluema (1; G. ivorensis; tropical West and Central Africa).

Sarraceniales Lindl. in C. F. P. von Martius, Consp. Regn. Veg.: 61. Sep-Oct 1835 [’Sarracenieae’]; Heliamphoraceae Chrtek, Slavíková et Studnička in Preslia 64: 8. 10 Feb 1992; Sarracenianae Thorne ex Reveal in Phytologia 79: 71. 29 Apr 1996; Sarraceniineae Reveal in Kew Bull. 66: 48. Mar 2011

Distribution Western United States, eastern North America, the Guayana Highlands.

Fossils Uncertain. Archaeamphora longiservia from the Lower Cretaceous of northeastern China may be a conifer.

Vegetative anatomy Mycorrhiza absent. Phellogen absent? Primary vascular tissue in scattered separate bundles. Secondary lateral growth absent. Vessel elements (in rhizome) with scalariform perforation plates; lateral pits scalariform, opposite or alternate, simple and/or bordered pits. Imperforate tracheary xylem elements tracheids with bordered pits, non-septate. Wood rays uniseriate or multiseriate, heterocellular?, or absent. Axial parenchyma apotracheal diffuse. Sieve tube plastids S type. Nodes? Crystals?

Leaves Alternate spiral, simple, entire, usually in basal rosette (in some species of Heliamphora on upright, usually unbranched stem), with ? ptyxis. Leaves ascidiate, infundibuliform or tubular, usually directed inwards (in Darlingtonia twisted 180° hence directed outwards), modified into water-holding (not in Heliamphora?) insect-trapping organs, usually with lid- or hood-like dorsal terminal lobe. Funnel-shaped zone of lamina in Heliamphora and Sarracenia consisting of adaxially fused margins (in Sarracenia with adaxial keel) and in uppermost part operculum (in Heliamphora small). In Darlingtonia a winged sheathing lower zone, a keeled tubular zone above and, in uppermost part, a stout dome-shaped almost closed zone. Distally, especially around entrance of pitcher, a nectar-secreting zone with nectariferous glands. Inner side of trap with a zone covered by downwardly directed hairs or bristles (in Darlingtonia scattered waxy platelets). In lower part of lamina an absorption zone. Proteolytic enzymes absent from pitcher leaf, nutrients instead becoming available to plant by bacterial decomposition of animal bodies in pitcher liquid. Lepidote or ensiform leaves sometimes forming late in season. Stipules and leaf sheath absent. Colleter-like glandular hairs present on adaxial side of petiole base. Petiole vascular bundle transection hippocrateromorphous (Ω-shaped), collateral, each bundle surrounded by sclerenchyma and starchy tissue. Venation pinnate? Stomata anomocytic. Cuticular wax crystalloids as platelets (in Darlingtonia, detachable, on inner side of pitcher). Leaf margin crenate to entire.

Inflorescence Flowers Sarracenia and Darlingtonia solitary, terminal; in Heliamphora usually terminal few-flowered raceme-like inflorescences. Floral prophylls (bracteoles) three.

Flowers Actinomorphic, often large. Hypogyny. Sepals (three or) four to six, with imbricate aestivation, persistent, free (in Heliamphora petaloid). Petals (three or) four to six, with imbricate aestivation, caducous, free (absent in Heliamphora). Ovary wall in Sarracenia with ten nectaries on stylar base above staminal groups; flowers in Darlingtonia and Heliamphora without nectaries. Disc absent.

Androecium Stamens eight to ten (Heliamphora) or numerous (Darlingtonia, Sarracenia), often in fascicles (in Sarracenia usually originating from ten staminal primordia). Filaments short, free or connate in ten fascicles (in Sarracenia), free from tepals. Anthers basifixed (Darlingtonia, Heliamphora) or dorsifixed (Sarracenia), non-versatile (Darlingtonia, Heliamphora) or versatile (Sarracenia), tetrasporangiate, introrse, usually longicidal (dehiscing by longitudinal slits; in Heliamphora poricidal, with short pore-like slits on caudate appendages). Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains (3–)4–5-colpor(oid)ate (Heliamphora), 4–6-colpor(oid)ate (Darlingtonia) or 6–9-colpor(oid)ate (Sarracenia), shed as monads, bicellular at dispersal. Exine tectate, with granular infratectum, verruculate (Heliamphora) or scrobiculate (Darlingtonia, Sarracenia).

Gynoecium Pistil composed of three (Heliamphora) or five (Darlingtonia, Sarracenia) connate carpels. Ovary superior, trilocular (Heliamphora) or quinquelocular (often incompletely divided in upper part). Style trilobate and apically truncate (Heliamphora), quinquelobate (Darlingtonia) or widened like an umbrella (Sarracenia), hollow?, in Sarracenia nectariferous. Stigmas three or five, small (in Sarracenia present on lower sides of five lobe apices), papillate, Dry type. Pistillodium absent.

Ovules Placentation axile in lower part, often intrusively parietal in upper part. Ovules numerous per carpel, anatropous, horizontal, usually unitegmic (in Darlingtonia bitegmic?), tenuinucellar. Integument approx. four cell layers thick. Hypostase? Megagametophyte monosporous, Polygonum type. Antipodal cells sometimes persistent. Endosperm development cellular. Endosperm haustoria? Embryogenesis caryophyllad.

Seeds Aril? Seed coat with wings or hairs. Exotesta with thick radial and inner cell walls, with theoid exotestal thickenings. Inner testal layers crushed. Perisperm not developed. Endosperm copious, oily and proteinaceous. Embryo straight, well differentiated, chlorophyll? Cotyledons two. Germination phanerocotylar.

Phytochemistry Flavonols (kaempferol, quercetin), O-methyl flavonols; cyanidin Route I iridoids (also decarboxylated iridoids), Group VI secoiridoids (morroniside), sarracenin (enoldiacetal monoterpene), and alkaloids present. sometimes present. Ellagic acid, saponins and cyanogenic compounds not found. Tannins?

Fossils Uncertain. Fossil wood (Sladenioxylum africanum), resembling the wood in extant Sladenia, has been found in Albian-Cenomanian layers in eastern Sudan.

Vegetative anatomy Phellogen ab initio pericyclic. Vessel elements in Sladenia arranged in radial groups. Vessel elements with scalariform perforation plates; lateral pits scalariform (Ficalhoa), opposite to alternate, simple or bordered pits. Imperforate tracheary xylem elements fibre tracheids with bordered pits, non-septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty vasicentric. Sieve tube plastids S type? Nodes 1:1?, unilacunar with one? leaf trace. Exudate in Ficalhoa white, latex-like. Chambered crystals present or absent. Calciumoxalate prismatic crystals or druses.

Leaves Alternate (spiral or distichous), simple, entire, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate; petiole in Sladenia also with wing bundles (associated fibres absent in Ficalhoa). Venation pinnate. Stomata anomocytic. Cuticular wax crystalloids? Epidermis in Sladenia with mucilage cells. Mesophyll with calciumoxalate druses. Sclereids absent. Leaf margin usually serrate (sometimes entire).

Flowers Actinomorphic, small. Hypogyny. Sepals four to seven, with imbricate aestivation, persistent, free (Sladenia) or connate (Ficalhoa). Petals five (or six), spiral or whorled, with imbricate aestivation, free (Sladenia) or connate at base (Ficalhoa). Spiral scale-like leaves present below perianth. Nectaries present on lower part of ovary (Sladenia). Disc absent.

Androecium Stamens (eight to) ten to 15, in one whorl. Filaments free, swollen (Sladenia) or connate in five fascicles, narrow (Ficalhoa), in Sladenia free from tepals, in Ficalhoa adnate to petals. Anthers basifixed, non-versatile, tetrasporangiate, introrse (Sladenia), poricidal (in Sladenia dehiscing by apical pores; in Ficalhoa by apical transverse slits). Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis successive. Pollen grains tricolp(or)ate (Sladenia), shed as monads, bicellular at dispersal. Exine tectate, with columellate? infratectum, verruculate or smooth.

Gynoecium Pistil composed of (two or) three (Sladenia) or five (or six) antepetalous (Ficalhoa) connate carpels. Ovary superior, in Sladenia usually trilocular (rarely bilocular), in Ficalhoa? usually quinquelocular (sometimes sexalocular). Style single, usually simple, short (sometimes with short acute lobes). Stigmas (two or) three (Sladenia), punctate, type? Pistillodium absent.

Ovules Placentation apical-axile (Sladenia) or axile (Ficalhoa). Ovules two collateral anatropous pendulous and epitropous (Sladenia) or numerous (Ficalhoa) per carpel, bitegmic, tenuinucellar. Micropyle endostomal (Sladenia). Outer integument three or four cell layers thick (Sladenia). Inner integument three or four cell layers thick (Sladenia). Obturator, hypostase, nucellar beak, and nucellar cap absent (Sladenia). Megagametophyte tetrasporous, 8-nucleate, Adoxa type. Endosperm development nuclear? Endosperm haustoria? Embryogenesis?

Fruit A loculicidal capsule with persistent calyx and central columella (Ficalhoa) or a schizocarp? with thin exocarp, crustaceous endocarp and three mericarps (Sladenia).

Seeds Aril absent. Seed coat in Sladenia winged. Testa crustaceous, with polygonal cells, little thickened (Ficalhoa). Perisperm not developed.. Endosperm copious. Embryo straight, well differentiated, chlorophyll? Cotyledons two, short. Germination?

Cytology n = 24 (Sladenia) – Sladenia has cell wall development of monocotyledonous type (also in Ficalhoa?).

de Candolle et Sprengel (transl. W. Jameson), Elem. Philos. Pl.: 140. Jul 1821 [’Styraceae’], nom. cons.

Halesiaceae D. Don in Edinburgh New Philos. J. 6: 49. Oct-Dec 1828; Halesiales Link, Handbuch 1: 667. 4-11 Jul 1829; Styracopsida Bartl., Ord. Nat. Plant.: 121, 158. Sep 1830 [’Styracinae’]; Styracales Rich. in C. F. P. von Martius, Consp. Regn. Veg.: 26. Sep-Oct 1835 [’Styraceae’]

Distribution The United States, Mexico, the West Indies, Central America, South America south to Argentina, the northern and eastern Mediterranean, eastern Himalayas (Nepal to Arunachal-Pradesh), Assam, eastern India, East Asia to the Korean Peninsula and Japan, Southeast Asia, Malesia, New Guinea and Solomon Islands.

Fossils Fossils assigned to Rehderodendron have been found in Eocene to Miocene layers of Europe.

Habit Usually bisexual (in Bruinsmia gynodioecious), evergreen or deciduous trees or shrubs.

Vegetative anatomy Phellogen ab initio pericyclic. Vessel elements usually with scalariform perforation plates (with numerous cross-bars; vessel elements rarely with simple perforation plates); lateral pits opposite or alternate (sometimes scalariform), bordered pits. Imperforate tracheary xylem elements fibre tracheids or libriform? fibres usually with bordered (sometimes also simple) pits, non-septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty. Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace. Secretory cavities with resinous canals often present. Silica present in wood at least in many species of Styrax. Prismatic calciumoxalate crystals present in some species.

Trichomes Hairs multicellular, uniseriate, stellate, peltate or lepidote (often brown to reddish-brown).

Leaves Alternate (spiral), simple, entire, with conduplicate-plicate or supervolute ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate or D-shaped; medullary bundles and/or wing bundles sometimes present (in Parastyrax complex). Venation pinnate, camptodromous. Stomata usually anomocytic. Cuticular wax crystalloids? Calciumoxalate as druses and/or solitary prismatic crystals. Leaf margin serrate, crenate or entire; leaf teeth gland-tipped.

Inflorescence Terminal or axillary, often raceme-like or paniculate (rarely one- or two-flowered) cymose. Floral prophylls (bracteoles) often absent.

Flowers Actinomorphic. Pedicel often articulated. Hypanthium present or absent. Half epigyny or sometimes (Halesia) epigyny. Sepals four or five (to nine), with open or valvate aestivation, persistent, connate. Petals (four or) five (to eight), with imbricate or subinduplicate-valvate aestivation, connate (in Halesia largely connate). Nectary absent. Disc absent.

Androecium Stamens (five to) eight to ten (to 20), usually twice as many as (rarely the same number or three or four times as many as) petals, usually obdiplostemonous (rarely haplostemonous, alternipetalous). Filaments stout (not distinctly delimited from anthers), usually flattened, usually connate below, usually adnate to petals (epipetalous). Anthers basifixed, non-versatile, tetrasporangiate, introrse or latrorse, longicidal (dehiscing by longitudinal slits); connective sometimes apically prolonged. Tapetum secretory. Female flowers with five staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolpor(oid)ate or tetracolpor(oid)ate, shed as monads, bicellular at dispersal. Exine tectate, with columellate infratectum, perforate, spinulate, verruculate or smooth.

Gynoecium Pistil composed of usually two to four (sometimes five) connate, alternisepalous, antepetalous carpels, often hairy inside; median carpel abaxial. Ovary inferior or semi-inferior, usually bilocular to quadrilocular in lower part (sometimes quinquelocular), usually unilocular in upper part (in Parastyrax multilocular). Style usually single, simple, filiform (occasionally lobed), hollow (with stylar canal). Stigma punctate, capitate or somewhat lobate, Dry type. Pistillodium absent.

Ovules Placentation usually axile (rarely subbasal). Ovules usually four to nine (rarely one or up to c. 30) per carpel, anatropous or hemianatropous, pendulous to ascending, usually apotropous (sometimes epitropous), unitegmic or bitegmic (Styrax), tenuinucellar. Micropyle endostomal. Outer integument in Styrax ? cell layers thick. Inner integument ? cell layers thick. Placental obturator present in some species. Megagametophyte monosporous, Polygonum type. Endosperm development cellular. Endosperm haustoria? Embryogenesis solanad.

Fruit A loculicidal capsule (Alniphyllum, Huodendron, Styrax), a drupe (Parastyrax, Styrax), a samara (Halesia, Pterostyrax), or a berry-like (Bruinsmia) or nutlike (Pterostyrax, Melliodendron, Rehderodendron, Sinojackia, Styrax) fruit with persistent hypanthium and calyx.

Seeds Aril? Seed coat in Alniphyllum winged. Testa thin, vascularized, crushed. Tegmen in Styrax? Perisperm not developed. Endosperm copious, oily. Embryo straight or somewhat curved, well differentiated, without chlorophyll. Cotyledons two, flat or almost terete. Germination phanerocotylar.

Phytochemistry Flavonols (kaempferol, quercetin), cyanidin, oleanolic acid derivatives, ellagic acid, tannins, triterpene saponins, caffeic acid, and hydroxycinnamic acids (p-coumaric acid, ferulic acid) present. Myricetin, alkaloids and cyanogenic compounds not found.

Use Ornamental plants, medicinal plants, balsam resins (benyamin-gum, benzoin etc. from Styrax) for pharmaceutical, confectionery and perfume industries, incense.

Desfontaines in Mém. Mus. Natl. Hist. Nat. 6: 9. 1820 [’Symplocearum’, ’Symploceae’], nom. cons.

Distribution Tropical and subtropical regions in southern, eastern and southeastern Asia, Malesia to New Guinea, eastern Australia, Melanesia, southeastern United States, southern Mexico, Central America, the West Indies, tropical South America; some species in temperate regions in East Asia and eastern North America.

Fossils Fossil Triporopollenites pollen from the Maastrichtian of California has been assigned to Symplocos. Fossil endocarps, pollen grains and leaves of Symplocaceae have been found in sediments from the Early Eocene to Miocene of Europe. Symplocos Subgenus Hopea as well as the extinct Pallioporia and Sphenotheca occur as fossil endocarps in European Eocene and later layers and Sphenotheca is very similar to some extant species of Symplocos in East Asia. During the Palaeogene Symplocos seems to have been widely distributed and largely dominating on the North American and Eurasiatic continents.

Habit Usually bisexual (rarely polygamomonoecious or dioecious), usually evergreen trees or shrubs (Symplocos paniculata is deciduous).

Vegetative anatomy Phellogen ab initio superficial. Vessel elements with scalariform perforation plates (often with numerous cross-bars); lateral pits scalariform, alternate or opposite, simple and/or bordered pits. Vestured pits sometimes present. Imperforate tracheary xylem elements tracheids with bordered pits, non-septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty, or absent. Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace. Calciumoxalate as crystal sand and lumps of prismatic crystals.

Trichomes Hairs unicellular or multicellular, uniseriate, septate (rarely with swollen base), or absent.

Leaves Alternate (usually spiral, sometimes distichous), simple, entire, often coriaceous, with supervolute ptyxis. Stipules and leaf sheath absent. Petiole vascular bundles? Venation pinnate. Stomata usually paracytic and as sparsely occurring large water stomata (protruding unicellular water-secreting, bladder-like on outer part, constricted internally where they run into lamina via rosette of epidermal cells). Cuticular wax crystalloids? Leaf margin serrate, glandular serrate (usually with caducous glands) or entire.

Inflorescence Usually axillary (rarely terminal), spike, raceme, panicle, fasciculate or head-like cymose (flowers sometimes solitary axillary).

Flowers Actinomorphic. Pedicel usually articulated (in Cordyloblaste clade not articulated). Usually epigyny (rarely half epigyny). Sepals (three to) five, with valvate or imbricate aestivation, persistent, connate at base. Petals (three to) five or ten (or eleven), with imbricate aestivation, deeply bifid, entirely fused into tube or connate only at base. Nectariferous disc annular, cylindrical or quinquelobate, with five glands.

Androecium Stamens usually numerous (sometimes three to five antesepalous, or ten), obdiplostemonous. Filaments fused into tube or connate only at base all or in five antesepalous alternipetalous fascicles (sometimes? free), usually adnate to corolla tube (epipetalous). Anthers almost spherical, basifixed to dorsifixed, versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Female flowers with staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains (2–)3(–4)-colporate, shed as monads, bicellular at dispersal. Exine tectate to semitectate, with granular or columellate infratectum, punctate, perforate to reticulate, psilate, scabrate, rugulate, areolate, verrucate, spinulate or echinate.

Gynoecium Pistil composed of two to five connate carpels; median carpel abaxial. Ovary usually inferior (rarely semi-inferior), usually trilocular (sometimes bi-, quadri- or quinquelocular), in upper part with incomplete septa, nectariferous. Style single, simple, narrow or wide, usually hollow (with stylar canal). Stigma small, capitate or bilobate to quinquelobate, Dry type or Wet type. Male flowers with pistillodium.

Ovules Placentation mostly axile. Ovules (two to) four per carpel, anatropous, pendulous, epitropous, unitegmic, tenuinucellar. Integument ? cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development cellular. Endosperm haustoria? Embryogenesis?

Fruit Usually a one-seeded drupe with persistent calyx. Pyrene with as many germination pores as fertile carpels.

Seeds Aril absent. Testa vascularized. Inner walls of exotestal cells usually thick (theoid exotestal thickenings) or thin. Endotesta? Perisperm not developed. Endosperm copious, oily and often starchy. Embryo large, straight or curved (sometimes U-shaped), chlorophyll? Cotyledons two, very short. Germination phanerocotylar.

DNA Mitochondrial intron coxII.i3 lost. Mitochondrial coxI intron present.

Phytochemistry Flavonols (kaempferol), O-methyl flavonols, cyanidin, catechins, davidigenin, Group II decarboxylated iridoids (e.g. cornine), oleanolic acid derivatives, arjunolic acid derivatives, ellagic and gallic acids, hydrolyzable tannins, proanthocyanidins (prodelphinidins), saponins, and lignans (pinoresinol) present. Indole alkaloids rare. Myricetin and cyanogenic compounds not found. Aluminium accumulated in many species.

Ternstroemiales Mirb. in C. F. P. von Martius, Consp. Regn. Veg.: 52. Sep-Oct 1835 [‘Ternstroemiaceae’]

Distribution Tropical and subtropical regions, the Canary Islands, tropical West and Central Africa, the Himalayas, East Asia, Southeast Asia, Malesia to New Guinea and Queensland, Melanesia, islands in southwestern Pacific, the Hawaiian Islands, Mexico, Central America, the West Indies, tropical South America.

Fossils Campylotropous seeds assigned to Eurya (e.g. Eurya crassitesta) and Visnea have been described from the Santonian onwards of Central Europe and from the Maastrichtian of Germany, and fossil pentamerous capsules and seeds of Protovisnea from the Late Turonian to the Maastrichtian. Seeds of Eurya having thick sclerotic testa are frequently recorded from Cenozoic sediments.

Vegetative anatomy Phellogen ab initio superficial. Medulla often septated by diaphragms consisting of sclereids. Narrow primary medullary rays alternating with wide ones. Vessel elements usually with scalariform (rarely reticulate) perforation plates; lateral pits scalariform or opposite (rarely alternate), simple and/or bordered pits. Vestured pits sometimes present. Imperforate tracheary xylem elements (tracheids? or) fibre tracheids with bordered pits, non-septate. Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse, or paratracheal scanty, or absent. Tyloses often abundant. Sieve tube plastids S type. Nodes usually 1:1, unilacunar with one leaf trace (rarely 3:3, trilacunar with three traces). Calciumoxalate as prismatic crystals.

Leaves Alternate (spiral or distichous), simple, entire, usually coriaceous, with conduplicate-involute or supervolute ptyxis. Stipules and leaf sheath absent. Colleters sometimes present (in, e.g., Freziera). Perulae absent. Petiole vascular bundles one to five, arcuately arranged. Venation usually brochidodromous (sometimes reticulodromous). Stomata usually anomocytic (sometimes cyclocytic). Cuticular waxes? Domatia in Eurya as pits. Epidermis in some species with mucilage cells. Mesophyll and petiolar cortex with sclerenchymatous idioblasts containing simple or branched sclereids. Leaf margin usually serrate (sometimes entire), with blackish caducous gland at leaf apex and tooth apices.

Inflorescence Flowers usually solitary axillary (sometimes in few-flowered axillary racemose inflorescence). Bracts with blackish gland at apex.

Flowers Actinomorphic. Usually hypogyny (rarely epigyny). Sepals (four or) five (to seven), with imbricate quincuncial aestivation, persistent, usually free (sometimes connate at base), usually with small gland at apex. Petals (four or) five (to seven; in Archboldiodendron usually 5+5), with imbricate quincuncial aestivation, usually free (sometimes connate at base). Nectaries present or absent.

Androecium Stamens (five to) numerous, often inflexed at apex. Filaments wide, flat or thick, often narrowing against apex, shorter than to at least twice as long as anthers, free or connate at base in five fascicles (in Visnea in antepetalous pairs), often adnate to petals at base. Anthers basifixed, non-versatile, tetrasporangiate, introrse, usually longicidal (dehiscing by longitudinal slits; sometimes poricidal, dehiscing with short pore-like slits; anthers in Eurya septate); connective usually prolonged, crystalliferous. Tapetum secretory, with binucleate to quadrinucleate cells. Female flowers usually with staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolporate, shed as monads, bicellular at dispersal. Exine tectate, with columellate? infratectum, psilate to scabrate, rugulate or foveolate.

Gynoecium Pistil composed of (two or) three to five (or six) connate carpels. Ovary usually superior (rarely inferior), (bilocular or) trilocular to quinquelocular (or sexalocular). Style single, hollow, simple or branched at apex, or stylodia several, free. Stigmas usually capitate?, papillate, Wet? type. Male flowers usually with pistillodium.

Ovules Placentation usually axile (rarely apical or parietal). Ovules (one to) numerous per carpel, anatropous (amphitropous?) or campylotropous, apotropous or epitropous, bitegmic, usually tenuinucellar. Micropyle endostomal. Outer integument three to nine cell layers thick. Inner integument approx. three cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis solanad.

Fruit In Freziereae usually a berry; in Ternstroemia an irregularly dehiscing capsule; in some species of Freziera a drupe; in Visnea a nut surrounded by fleshy accrescent calyx.

Seeds Aril absent. Seed coat mesotestal. Sarcotesta present in Anneslea and Ternstroemia. Exotestal cells often enlarged, sometimes with thickened walls, with theoid exotestal thickenings. Mesotesta of various thickness, often with lignified cell walls, sometimes with sclereids, often crystalliferous. Endotesta? Tegmen? Perisperm not developed. Endosperm copious or sparse, oily. Embryo large, U-shaped to almost straight, well differentiated, without chlorophyll. Cotyledons two, incumbent, shorter than radicula. Germination phanerocotylar?

Cytology n = 20, 25 (Ternstroemieae); n = 12, 15, 18, 21–23 or more (21 most frequent) (Freziereae)

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), cyanidin, triterpene acids (betulinic acid), ellagic acid, proanthocyanidins (prodelphinidins), and saponins present. Iridoids not found. Aluminium accumulated in many species.

Pellicieraceae (Triana et Planch.) L. Beauvis. ex Bullock in Taxon 8: 192. 24 Jul 1959

Distribution West Malesia, southern Venezuela, mangroves along the Pacific coast and islands in Central America, Colombia and northern Ecuador, and the Atlantic coast in Colombia.

Fossils Fossil pollen grains (Lanagiopollis crassa) assigned to Pelliciera from the Eocene onwards indicate that during the Paleogene this group was widely distributed along the Caribbian and Central American coasts, and also along the Nigerian coast.

Habit Bisexual, evergreen trees or shrubs (Pelliciera consists of mangrove trees with buttresses). Aerial roots frequent along stem base in Pelliciera.

Vegetative anatomy Phellogen ab initio inner-cortical (Pentamerista, Tetramerista) or superficial (Pelliciera). Vessel elements usually with simple (sometimes scalariform) perforation plates; lateral pits usually alternate (rarely opposite), bordered pits. Imperforate tracheary xylem elements fibre tracheids or libriform fibres with simple or bordered pits, septate or non-septate. Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty (in Tetramerista also ray-adjacent). Sieve tube plastids S type? Nodes 3:3, trilacunar (in Pelliciera unilacunar?) with three leaf traces. Cortex and medulla in Pelliciera with branched sclerenchymatous idioblasts and thick-walled fibres. Heartwood in Tetramerista with gum-like substances. Parenchyma in Pelliciera with calciumoxalate raphides in raphid sacs. Crystal sand present or absent.

Leaves Alternate (spiral), simple, entire, coriaceous (in Pelliciera somewhat asymmetrical), with supervolute (Tetramerista, Pentamerista) or involute (Pelliciera) ptyxis. Stipules and leaf sheath absent. Petiole in Pelliciera with one pair of extrafloral nectaries. Petiole vascular bundle transection in Pelliciera annular. Venation pinnate. Stomata usually anomocytic (sometimes paracytic; in Pelliciera cyclocytic). Cuticular wax crystalloids? Abaxial side of lamina in Tetramerista with black glandular dots, in Pentamerista with domatia along veins. Mesophyll in Tetramerista with sclerenchymatous idioblasts with calciumoxalate raphides and often crystal sand. Spongy tissue in Pelliciera with stone cells. Margin of young leaves glandular-serrate, with gland-like structures (in Pelliciera salt-excreting?).

Inflorescence In Pentamerista and Tetramerista axillary, umbel-like, corymbose or capitate cymose; flowers in Pelliciera solitary, axillary. Bracts in Pelliciera large and petaloid. Bud scales absent in Pelliciera. Floral prophylls (bracteoles) often large, caducous or persistent.

Flowers Actinomorphic, usually small (in Pelliciera large). Hypogyny. Sepals four or five, with imbricate aestivation, petaloid, unequal in size, with shining glandular dots on adaxial side near middle (Tetramerista) or close to base? (Pelliciera; absent in Pentamerista), persistent or caducous, free. Petals four or five, with imbricate aestivation, in Tetramerista and Pelliciera with adaxial glandular dots, persistent or caducous, free. Nectaries as bottle-shaped glands at calyx base. Disc absent.

Androecium Stamens four or five, haplostemonous, antesepalous, alternipetalous. Filaments flattened, inflexed in bud (finally reversed), at base usually somewhat connate (in Pelliciera free), free from tepals. Anthers in Pentamerista and Tetramerista dorsifixed, in Pelliciera basifixed and sagittate with apical appendage, non-versatile, tetrasporangiate, usually introrse (finally extrorse; in Pelliciera extrorse to latrorse), longicidal (dehiscing by longitudinal slits); connective in Pelliciera apically prolonged. Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolporate (constricticolporate), shed as monads, bicellular at dispersal. Exine semitectate, with granular infratectum, reticulate (in Pelliciera also scabrate).

Gynoecium Pistil composed of usually four or five (in Pelliciera two) connate carpels. Ovary superior, quadrilocular or quinquelocular (in Pelliciera bilocular, one locule sterile or aborted, although basically quinquecarpellate). Style single, simple (in Pelliciera persistent in fruit). Stigma punctate or somewhat lobate (in Pelliciera bifid), type? Pistillodium absent.

Ovules Placentation subapical (Pelliciera) to basal (Tetramerista, Pentamerista). Ovule one per carpel, in Pentamerista and Tetramerista anatropous and ascending (epitropous?), in Pelliciera campylotropous and pendulous, (incompletely) bitegmic, incompletely tenuinucellar. Micropyle endostomal? Outer integument ? cell layers thick. Inner integument ? cell layers thick. Megagametophyte monosporous, Polygonum type? Endosperm development nuclear? Endosperm haustorium micropylar? Embryogenesis?

Fruit A four- or five-seeded coriaceous berry (in Pelliciera up to 13 cm in diameter, spongy inside and with persistent style).

Seeds Aril absent. Testa in Tetramerista and Pentamerista multi-layered; testa in Pelliciera absent, with mature seed consisting of large naked embryo, hemispherical cotyledons surrounding red plumule, and radicula at maturation penetrating stylar beak of fruit. Tegmen? Perisperm not developed. Endosperm usually copious (absent? in Pelliciera). Embryo large, straight, well differentiated, chlorophyll? Cotyledons two, usually much shorter than hypocotyl (in Pelliciera large, fleshy). Germination phanerocotylar.

Camelliaceae Mirb. ex A. P. de Candolle, Essai Propr. Méd. Pl., ed. 2: 97. 11 Mai 1816 [‘Camellieae’]; Theales Bercht. et J. Presl, Přir. Rostlin: 219. Jan-Apr 1820 [‘Theaceae’]; Gordoniaceae (DC.) Spreng., Syst. Veg. 3: 12. Jan-Mar 1826; Camelliales Link, Handbuch 2: 346. 4-11 Jul 1829 [’Camelliaceae’]; Gordoniales J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 180. 1846 [‘Gordoniaceae’]; Malachodendraceae J. Agardh, Theoria Syst. Plant.: 130. Apr-Sep 1858 [’Malachodendreae’], nom. illeg.; Theineae Engl., Syllabus, ed. 2: 150. Mai 1898

Distribution Southeastern United States, the West Indies, Central America, tropical South America, East Asia to the Korean Peninsula and Japan, Southeast Asia, Malesia, New Guinea.

Fossils A large number of seeds from the Late Cretaceous and especially from the Cenozoic have been assigned to Theaceae, although it is sometimes doubtful whether these should be attributed to Theaceae. Palaeoschima consists of fruits and seeds that has been described from the Late Cretaceous, but the systematic affiliation is problematic. Camellia is represented by a number of Cenozoic fossils and was formerly distributed also in North America. Fossil flowers of Theaceae have been found in amber from the Baltic area, and several extant genera are known from the Cenozoic of Europe.

Vegetative anatomy Phellogen ab initio usually close to pericycle (rarely subepidermal). Pericyclic envelope collenchymatous, lignified or with groups of fibres. Vessel elements with scalariform perforation plates; lateral pits scalariform, alternate or opposite, simple pits. Vestured pits sometimes present. Imperforate tracheary xylem elements tracheids or fibre tracheids with bordered pits, non-septate. Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty, or absent. Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace. Sclereids abundant. Mucilage cells frequently present. Calciumoxalate as prismatic crystals.

Trichomes Hairs unicellular or multicellular, usually uniseriate (sometimes stellate, rarely fasciculate), or absent.

Leaves Alternate (usually spiral, sometimes distichous), simple, entire, usually coriaceous, usually with involute or supervolute (sometimes conduplicate) ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate (U- or V-shaped), sometimes also with lateral bundles. Venation pinnate, craspedodromous. Stomata usually paracytic, anisocytic to cyclocytic, gordoniaceous type, with guard cells surrounded by two to four narrow subsidiary cells (rarely anomocytic). Cuticular wax crystalloids as rodlets. Epidermal cells sometimes mucilaginous. Mesophyll and petiolar cortex with or without sclerenchymatous idioblasts containing numerous branched thickened sclereids (often visible as points or dots). Leaf margin usually serrate (rarely entire), with small caducous gland associated with each tooth (theoid teeth).

Inflorescence Flowers axillary, solitary. Buds usually perulate. Floral prophylls (bracteoles) two or more, alternate (spiral) and grading into usually whorled (sometimes spiral) sepals.

Flowers Actinomorphic, often large. Usually hypogyny (rarely half epigyny). Sepals (four or) five to seven, with imbricate aestivation, often persistent, usually free or connate at base. Petals usually five (sometimes 5+5, rarely numerous), with imbricate aestivation, free or connate at base, sometimes spiral. Nectary absent or nectar produced at base of filaments or ovary. Disc absent.

Androecium Stamens (ten to) 20 to c. 40 (to much more than 100), in two to five whorls, produced from five antepetalous primordia or an annular primordium. Filaments usually free (sometimes in five antepetalous fascicles; rarely connate into a tube), often adnate to petal bases. Anthers articulated, usually dorsifixed (rarely basifixed), often versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits); connective sometimes prolonged. Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolpor(oid)ate, shed as monads, bicellular at dispersal. Exine tectate, with columellate infratectum, rugulate or perforate to finely reticulate, scabrate. Pseudopollen formed in cells of connective.

Gynoecium Pistil composed of (three to) five (to ten) connate, usually antepetalous carpels. Ovary usually superior (rarely semi-inferior), (trilocular to) quinquelocular (to decemlocular), often nectariferous. Style single, simple or branched, or stylodia several, free. Stigma capitate or slightly lobate or stigmas punctate, papillate, Wet type. Pistillodium absent.

Ovules Placentation usually axile (sometimes basal). Ovules two or more per carpel, usually anatropous (in Schima campylotropous), usually pendulous, apotropous or pleurotropous, bitegmic, tenuinucellar. Micropyle endostomal. Outer integument four to ten cell layers thick. Inner integument four to eleven cell layers thick. Hypostase present. Megagametophyte usually monosporous, Polygonum type (in ‘Camellia’ disporous, Allium type). Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis solanad. Polyembryony occurring.

Fruit Usually a loculicidal (in Franklinia also septicidal) capsule often with persistent columella and calyx (in some species of Pyrenaria a drupe; a capsule in some species of ‘Camellia’ irregularly dehiscing).

Seeds Aril absent. Seed coat mesotestal, in Gordonia, Schima and Stewartia winged. Testa and tegmen vascularized. Exotesta with thickenings on radial and inner tangential cell walls (not on outer tangential walls), with theoid exotestal thickenings, often lignified. Mesotesta lignified (sometimes with fibres, sometimes with sclereids). Endotesta often lignified. Tegmen? Perisperm not developed. Endosperm usually sparse, oily, or absent. Embryo usually straight (rarely curved), without chlorophyll. Cotyledons two, large (longer than radicula). Germination phanerocotylar or cryptocotylar.

DNA Plastid gene rpl22 absent in Stewartia (present in nuclear genome?). Mitochondrial intron coxII.i3 lost.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), biflavonoids, catechins (e.g. epigallocatechin-3-gallate), cyanidin, oleanolic acid derivatives, triterpenes, gallic acid (in cultivated species of ‘Camellia’ Section Thea), ellagic acid, methylated ellagic acids, ellagitannins (pedunculagin, tellimagrandin II), proanthocyanidins (prodelphinidins), chlorogenic acid, alkaloids (purine bases such as theobromine, theofylline and caffeine in cultivated species of ‘Camellia’ Section Thea), and triterpene saponins present. Cyanogenic compounds not found. Aluminium and fluorides accumulated.

Use Ornamental plants, stimulants (Camellia sinensis), seed-oils (Camellia sasanqua), timber.

Don in Edwards’s Bot. Reg. 21: ad t. 1764. 1835 [’Theophrasteae’], nom. cons.

Distribution Southern Florida, Mexico, Central America, the West Indies, South America south to northern Paraguay.

Habit Usually bisexual (in Clavija usually dioecious, gynodioecious? or polygamodioecious), usually evergreen (rarely deciduous) trees or shrubs, often pachycaul.

Vegetative anatomy Phellogen? Primary medullary rays wide. Vessel elements with simple perforation plates; lateral pits alternate, bordered pits. Imperforate tracheary xylem elements libriform fibres with simple pits, septate or non-septate. Wood rays usually multiseriate, heterocellular. Axial parenchyma paratracheal scanty vasicentric, or absent. Sieve tube plastids S type? Nodes 1:1, unilacunar with one leaf trace, 3:3, trilacunar with three traces, or 5:5, pentalacunar with five traces. Subepidermal fibres sometimes non-lignified. Resins absent. Wood ray cells often with silica bodies and/or prismatic calciumoxalate crystals. Styloids present in some species.

Trichomes Hairs simple or branched; glandular hairs (on leaves) consisting of unicellular stalk and multicellular head; glandular hairs usually sunken.

Leaves Alternate (usually pseudoverticillate on branch tips), simple, entire, coriaceous, gland-dotted (often scale-like; in Jacquinia sometimes needle-like, in Theophrasta sometimes very large), with conduplicate (or involute?) ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection deeply arcuate or annular; petiole with small adaxial inverted bundles. Venation pinnate. Stomata? Cuticular wax crystalloids? Lamina on both sides usually striated by subepidermal fibres. Mesophyll with calciumoxalate druses. Leaf margin serrate (often serrate-dentate) or entire.

Inflorescence Usually terminal (in Clavija and Neomezia lateral), racemes, corymbs or panicles (flowers rarely solitary). Bracts inserted on pedicels. Floral prophylls (bracteoles) absent?

Flowers Actinomorphic. Hypogyny to half epigyny. Sepals (four or) five, with imbricate aestivation, persistent, free or somewhat connate at base. Petals (four or) five, with imbricate aestivation, usually slightly unequal in size, usually connate into a campanulate or infundibuliform corolla. Petals and stamens developing from common primordia. Nectary? Disc absent.

Androecium Stamens (four or) five, antepetalous, alternisepalous. Filaments flattened, connate usually only at base (in Clavija often entirely connate into tube), adnate to lower part of corolla tube. Anthers forming cone in centre of young flower, later spreading, basifixed to dorsifixed, sometimes versatile?, tetrasporangiate, extrorse or introrse, longicidal (dehiscing by longitudinal slits); connective in Neomezia and Theophrasta prolonged at apex, ab initio incurved above stigma. Nectariferous hairs sometimes present. Tapetum secretory, with uninucleate cells. Staminodia (four or) five, antesepalous, alternipetalous, extrastaminal, epipetalous (inserted above fertile stamens), petaloid, gibbous, narrow or triangular, often glanduliferous. Upper and lower parts of thecae filled with druses and single crystals of calciumoxalate.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolporate (or tetracolporate), shed as monads, bicellular at dispersal. Exine tectate or semitectate, with columellate infratectum, reticulate, foveolate, fossulate or rugulate.

Gynoecium Pistil composed of (four or) five connate antepetalous carpels. Ovary superior or semi-inferior, unilocular, mucilaginous. Style single, simple. Stigma capitate, punctate, discoid, truncate or somewhat lobate, non-papillate Dry type, or papillate Wet type. Pistillodium?

Ovules Placentation free central to basal (central column sometimes reduced). Ovules numerous per ovary, anatropous, hemitropous or campylotropous, ascending, bitegmic, tenuinucellar. Micropyle bistomal. Outer integument two to four cell layers thick. Inner integument ? cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit Usually a one- or several-seeded berry with dry (sometimes woody) pericarp and juicy placentae (rarely a one-seeded drupe).

Seeds Aril absent. Seed coat exotestal. Testa multiplicative. Exotestal cells flattened, with thick walls. Hypodermal cells sometimes with thick anticlinal walls. Mesotestal cells often with crystals. Endotesta? Tegmen? Perisperm not developed. Endosperm copious, hard, oily (in many species with thick cell walls with pores), amyloid? Embryo large, straight, well differentiated, chlorophyll? Cotyledons two, usually foliaceous (sometimes small). Germination phanerocotylar.

Phytochemistry Insufficiently known. Saponins present. Ellagic acid, proanthocyanidins, alkaloids, and cyanogenic compounds not found. Polyphenols? Glycosides?

Use Ornamental plants, fruits (Clavija), fish poison, soap (bark and cortex of Jacquinia).

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“ERICIDAE”

ERICALES Bercht. et J. Presl

ACTINIDIACEAE Gilg et Werderm.

BALSAMINACEAE A. Rich.

CLETHRACEAE Klotzsch

CYRILLACEAE (Torr. et A. Gray) Lindl.

DIAPENSIACEAE (Link) Lindl.

EBENACEAE Gürcke

ERICACEAE Juss.

FOUQUIERIACEAE DC.

LECYTHIDACEAE A. Rich.

MAESACEAE (A. DC.) Anderb., B. Ståhl et Källersjö

MARCGRAVIACEAE Bercht. et J. Presl

MITRASTEMONACEAE Makino

MYRSINACEAE R. Br.

PENTAPHYLACACEAE Engler

POLEMONIACEAE Juss.

PRIMULACEAE Batsch ex Borkh.

RORIDULACEAE Augier ex Martinov

SAMOLACEAE Raf.

SAPOTACEAE Juss.

SARRACENIACEAE Dumort.

SLADENIACEAE (Gilg et Werderm.) Airy Shaw

STYRACACEAE DC. et Spreng.

SYMPLOCACEAE Desf.

TERNSTROEMIACEAE Mirb. ex DC.

TETRAMERISTACEAE (H. Hallier) Hutch.

THEACEAE Mirb. ex Ker Gawl.

THEOPHRASTACEAE (Bartl.) G. Don