CELASTRALES Baskerville Main Tree, Synapomorphies

Fossils Uncertain. Wuyunanthus hexapetalus, a hexamerous flower from the Paleocene of northeastern China, has been assigned to Celastraceae, although its systematic affiliation is problematic.

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

Vegetative anatomy Phellogen ab initio usually subepidermal (sometimes epidermal or cortical). Secondary lateral growth normal or anomalous (from concentric cambia). Vessel elements usually with simple (rarely scalariform) perforation plates; lateral pits alternate or scalariform, simple or bordered pits. Imperforate tracheary xylem elements tracheids, fibre tracheids or libriform fibres with simple or bordered pits, septate or non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal, diffuse or diffuse in aggregates, or paratracheal scanty vasicentric, reticulate, unilateral, or banded (rarely aliform, lozenge-aliform, winged-aliform, or confluent; sometimes absent). Intraxylary phloem (concentric or diffuse) sometimes present. Sieve tube plastids S type. Nodes usually 1:1, unilacunar with one leaf trace (rarely 3:3, trilacunar with three traces, or 1:5–7, unilacunar with five to seven traces). Usually with laticifers containing gutta between cortex and phloem. Cortex with or without cristarque cells. Calciumoxalate druses sometimes present; prismatic crystals often abundant; solitary and aggregates of crystals rarely present.

Trichomes Hairs usually absent (rarely unicellular, simple, furcate or stellate); glandular hairs rarely present.

Leaves Usually alternate (spiral or distichous) or opposite (rarely verticillate), usually simple rarely palmately compound and unifoliolate, entire, usually with involute (rarely conduplicate) ptyxis. Stipules usually small, cauline and caducous; leaf sheath absent. Petiole vascular bundle transection arcuate or annular. Venation usually pinnate (rarely campylodromous or acrodromous, palmate). Stomata usually anomocytic or anisocytic (rarely laterocytic, cyclocytic, tetracytic, or paracytic). Cuticular wax crystalloids usually absent (rarely as platelets). Domatia as pockets or absent. Epidermis with or without mucilaginous idioblasts; epidermal cells sometimes with calciumoxalate as druses or rhomboidal crystals. Mesophyll with or without secretory cavities, with or without sclerenchymatous idioblasts. Laticifers with gutta present in association with vascular strands and sometimes in mesophyll. Leaf margin entire, serrate, glandular-serrate or spinose-serrate.

Inflorescence Terminal or axillary, cymose, compound thyrsoid, fasciculate, paniculate, umbellate, raceme-like or spicate, or flowers solitary.

Flowers Usually actinomorphic (rarely zygomorphic). Usually hypogyny (rarely epigyny, with stamens at ovary apex, or half epigyny with hypanthium surrounded by nectary). Sepals (two to) four or five (or six), usually with imbricate quincuncial (rarely valvate or open) aestivation, free. Petals (two to) four or five (or six), usually with imbricate quincuncial (rarely contorted or valvate) aestivation, persistent or caducous, usually free. Nectariferous disc annular, intrastaminal to extrastaminal (rarely absent), or nectaries intrastaminal at bases of staminodia. Floral tissues often with calciumoxalate druses.

Androecium Stamens usually three to five (rarely two, 5+5, or c. 30 to more than 100). Filaments usually free from each other (rarely connate at base into tube), free from tepals. Anthers basifixed to dorsifixed (rarely ventrifixed), usually non-versatile, tetrasporangiate or disporangiate, usually introrse to latrorse (sometimes extrorse), usually longicidal (usually dehiscing by longitudinal slits; rarely with transverse or oblique orifice, or with apical confluent horizontal slits). Tapetum secretory. Staminodia (two or) three to five, intrastaminal, or absent; female flowers with or without staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolpor(oid)ate (rarely dicolporate, tetracolporate, triporate or syncolpate), shed as monads or tetrads (rarely polyads with eight or sixteen pollen grains), bicellular or tricellular at dispersal. Exine usually semitectate (rarely tectate), with columellate infratectum, usually reticulate (rarely psilate or microreticulate).

Gynoecium Pistil composed of usually two to five (rarely more than five) connate antepetalous carpels (when three then median carpel adaxial; rarely three paracarp and connate carpels); carpels basically synascidiate; postgenital carpel closure through much elongated cells. Ovary usually superior (rarely inferior or semi-inferior), partially or entirely bilocular to quinquelocular (all locules except one usually degenerating; rarely primarily unilocular, or with eight to twelve locules each locule secondarily transversely divided by secondary septa). Style usually single, simple (stylodia sometimes two to five, free or connate at base), or absent. Stigmas two to five or one, capitate or lobate, commissural, non-papillate, Dry type. Male flowers with or without pistillodium.

Ovules Placentation usually axile (rarely parietal, basal or apical). Ovules one to twelve (to more than 50) per carpel, usually anatropous (rarely amphitropous), ascending to pendulous, usually apotropous (rarely epitropous), bitegmic, usually weakly crassinucellar (sometimes incompletely tenuinucellar). Micropyle usually bistomal (rarely endostomal or exostomal). Megagametophyte monosporous, Polygonum type. Synergids sometimes with a filiform apparatus. Antipodal cells usually ephemeral. Endosperm development usually nuclear (rarely cellular). Endosperm haustoria absent. Embryogenesis caryophyllad or solanad (sometimes asterad).

Fruit A loculicidal capsule, a berry, drupe or samara (rarely a septicidal or loculicidal-septicidal capsule, nut, or schizocarp with two to five nutlike mericarps).

Seeds Seed with or without funicular aril or arilloid structure from hilum or exostoma (sometimes winged; wings in at least some cases possibly homologous to an aril). Seed coat usually exotegmic (occasionally reduced). Testa multiplicative, usually lignified (sometimes vascularized). Exotesta with thick cuticle, often palisade (rarely tanniniferous and with thickened outer wall). Mesotesta with sclerotic cells (rarely absent or fibrous). Endotesta unspecialized? Exotegmen usually fibrous (consisting of tall lignified fibres). Endotegmen rarely persistent, tanniniferous. Perisperm not developed. Endosperm copious to sparse, oily, or absent. Embryo straight, well differentiated, usually with chlorophyll. Cotyledons two. Germination phanerocotylar or cryptocotylar.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), cyanidin, Ouratea catechins, diterpene and triterpene derivatives (celastrol, pristimerin, tingenin), gutta (trans-1,4-polyisoprene), tannins, proanthocyanidins (prodelphinidins), benzylisoquinoline alkaloids, pyrrolizidine alkaloids as aliphatic monocarboxylic esters, sesquiterpene alkaloids (i.a. mayteine and 7-epi-mayteine, euonymine and 7-epi-euonymine, mekongensines, peritassines), d-norpseudoephedrine (cathine), saponins, narcotic L(S)-(-)-α-aminopropiophenone (cathinone), maytansine (ansamycin macrolide), hexitols (dulcitol etc.), and salacinol present. Ellagic acid and cyanogenic compounds not found.

Celastraceae are sister to Parnassiaceae with very high support. They share the following potential synapomorphies (Stevens 2001 onwards): nodes 1:1; leaves with veins proceeding to congested deciduous tooth; stamens as many as petals; carpels antepetalous; stigma commissural; ovules tenuinucellar; presence of flavonols and hexitols (dulcitol etc.); and absence of ellagic acid. The leaf traces in both Brexia and Parnassia arise in the central part of the stem at a significant distant below the petiole.

Brown in M. Flinders, Voy. Terra Austral. 2: 554. 19 Jul 1814 [‘Celastrinae’], nom. cons.

Hippocrateaceae Juss. in Ann. Mus. Natl. Hist. Nat. 18: 486. Jul-Aug 1811 [‘Hippocraticeae’], nom. cons.; Stackhousiaceae R. Br. in M. Flinders, Voy. Terra Austr. 2: 555. 19 Jul 1814 [‘Stackhouseae’], nom. cons.; Euonymales Bercht. et J. Presl, Přir. Rostlin: 226. Jan-Apr 1820 [‘Evonymeae’] Hippocrateales Juss. ex Bercht. et J. Presl, Přir. Rostlin: 225. Jan-Apr 1820 [‘Hippocrateae’]; Euonymaceae Juss. ex Bercht. et J. Presl, Přir. Rostlin: 2(110): 438, 439. 1825 [‘Evonymeae’]; Brexiaceae Loudon, Hort. Brit.: 524. 30 Aug 1830 [‘Brexieae’]; Brexiales Lindl., Nix. Plant.: 18. 17 Sep 1833; Salaciaceae Raf., Fl. Tellur. 4: 101. med 1838 [‘Salacides’]; Celastropsida Brongn., Enum. Plant. Mus. Paris: xxiv, 87. 12 Aug 1843 [’Celastoideae’]; Chingithamnaceae Hand.-Mazz. in Sinensia 2: 126. Apr-Mai 1932; Stackhousiales R. Br. in C. F. P. von Martius, Consp. Regn. Veg.: 55. Sep-Oct 1835 [’Stackhousieae’]; Siphonodontaceae (Croizat) Gagnep. et Tardieu ex Tardieu in Notul. Syst. (Paris) 14: 102. Jul-Sep 1951, nom. cons.; Canotiaceae Airy Shaw in Kew Bull. 18: 255. 8 Dec 1965; Plagiopteraceae Airy Shaw in Kew Bull. 18: 266. 8 Dec 1965; Celastranae Takht., Sist. Filog. Cvetk. Rast. [Syst. Phylog. Magnolioph.]: 371. 4 Feb 1967; Pottingeriaceae (Engl.) Takht., Sist. Magnoliof. [Systema Magnoliophytorum]: 212. 24 Jun 1987

Distribution Mainly tropical and subtropical regions in the Northern and Southern Hemispheres; some species in temperate areas.

Habit Bisexual, monoecious, andromonoecious, polygamomonoecious?, dioecious, gynodioecious, or polygamodioecious?, usually evergreen or deciduous trees, shrubs or lianas (rarely herbs, suffrutices, or ericoid or epiphytic shrubs; Stackhousia usually annual or perennial herbs, sometimes succulent). With or without spines (Acanthothamnus and Canotia with glandular stems). Branches rarely photosynthesizing phyllocladia. Bark often yellow (due to triterpenic compounds).

Vegetative anatomy Phellogen ab initio usually subepidermal (sometimes epidermal or cortical). Medulla homogeneous or heterogeneous. Young stem with vascular cylinder. Secondary lateral growth normal or anomalous (via concentric cambia). Vessel elements usually with simple (rarely scalariform) perforation plates; lateral pits alternate, usually bordered (in, e.g., Hippocratea and Tripterygium sometimes also simple) pits. Imperforate tracheary xylem elements tracheids, fibre tracheids or libriform fibres with simple and/or bordered pits, septate or non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, heterocellular. Axial parenchyma apotracheal diffuse, or paratracheal scanty, reticulate, vasicentric, unilateral, or banded (rarely aliform, lozenge-aliform, winged-aliform, or confluent; sometimes, e.g. in Pottingeria, absent). Tyloses sometimes frequent (sometimes sclerotic). Intraxylary phloem (concentric or diffuse) present in Salacioideae. Sieve tube plastids S type. Endodermal cells in Stackhousia thick-walled and with granular content (endodermis absent in Pottingeria). Nodes usually 1:1, unilacunar with one leaf trace (in Brexia 3:3, trilacunar with three traces; in Lophopetalum 1:5–7, unilacunar with five to seven traces). Usually with latex sacs or laticifers containing gutta between cortex and phloem. Phloem in Pottingeria with sclereids. Cortex with or without cristarque cells. Colleters often blackening. Calciumoxalate druses present in flowers of some species. Prismatic crystals often abundant. Solitary and aggregated crystals present in cortex and phloem of Pottingeria.

Trichomes Hairs usually absent (rarely unicellular or multicellular, simple or furcate; in Plagiopteron stellate, ferrugineous); glandular hairs present in Stackhousioideae.

Leaves Usually alternate (spiral or distichous) or opposite (rarely verticillate), simple, entire, sometimes coriaceous, rarely reduced and scale-like, usually with involute (in Brexia and Plagiopteron conduplicate to flat) ptyxis. Stipules usually small, cauline and caducous (sometimes fringed or absent; sometimes persistent); colleters often present; leaf sheath absent. Petiole vascular bundle transection usually arcuate or annular (sometimes complex with several bundles). Venation usually pinnate (rarely acrodromous, palmate); lateral veins often proceeding into congested caducous foliar teeth. Stomata usually anomocytic or anisocytic (rarely laterocytic, cyclocytic or tetracytic). Cuticular wax crystalloids usually absent (rarely as platelets). Domatia as pockets or absent. Epidermis with or without mucilaginous idioblasts. Epidermal cells in some genera with calciumoxalate as druses or rhomboidal crystals. Mesophyll with or without secretory cavities, with or without sclerenchymatic idioblasts. Laticifers with gutta in association with vascular strands and sometimes in mesophyll (often visible as elastic threads when leaf cut). Leaf margin entire, serrate, glandular-serrate, or spinose-serrate.

Inflorescence Terminal or axillary (in ‘Polycardia’ sometimes epiphyllous), compound thyrsoid, fascicle, panicle, umbellate, raceme-like or spicate cymose, or flowers solitary.

Flowers Usually actinomorphic (rarely zygomorphic), usually small. Usually hypogyny (rarely epigyny, with stamens at ovary apex; in Stackhousioideae half epigyny with hypanthium surrounded by nectary; in Pottingeria half epigyny). Perianth in Plagiopteron with epicalyx. Sepals (two to) four or five (or six), usually with imbricate (rarely valvate; in Plagiopteron open) aestivation, free. Petals (two to) four or five (or six), usually with imbricate (rarely contorted or valvate) aestivation, persistent or caducous, usually free (in Stackhousia clawed; absent in Plagiopteron). Nectariferous disc usually wide, massive and often fleshy, annular (in Brexia with stiff outgrowths), intrastaminal to extrastaminal (absent in Plagiopteron). Floral tissues often with calciumoxalate druses.

Androecium Stamens (two or) three to five (in Stackhousioideae three longer and two shorter; in Plagiopteron c. 30 to more than 100), antesepalous, alternipetalous. Filaments usually free (in Plagiopteron connate at base into infundibuliform tube), free from tepals (in Pottingeria subulate, inserted at disc). Anthers basifixed or dorsifixed, sometimes versatile, tetrasporangiate or disporangiate, usually introrse to latrorse (in Hippocrateoideae usually extrorse), usually longicidal (usually dehiscing by longitudinal slits; rarely with transverse or oblique orifice; in Plagiopteron with apical confluent horizontal slits). Tapetum secretory, with binucleate (in, e.g., Stackhousia) to multinucleate cells. Staminodia (two or) three to five, intrastaminal, alternating with stamens, or absent (staminodia in Brexia fringed; in Pottingeria absent); female flowers with or without staminodia (stamens in Siphonodon alternating with staminodia).

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolporate (rarely dicolporate, tetracolporate or triporate), shed as monads or tetrads (rarely as polyads of eight or sixteen pollen grains), bicellular or tricellular at dispersal. Exine usually semitectate (in Plagiopteron tectate?), with columellate infratectum, usually reticulate (in Plagiopteron psilate; in Pottingeria microreticulate or reticulate), usually with endexinal folds in aperture.

Gynoecium Pistil composed of usually two to five (rarely more than five) connate antepetalous carpels (when three then median carpel adaxial; in Siphonodon numerous; in Pottingeria three paracarp and connate carpels); carpels sunken into disc. Ovary usually superior (rarely inferior or semi-inferior), partially or entirely usually bilocular to quinquelocular (all locules except one usually degenerating; rarely, e.g. in Empleuridium and Pottingeria, primarily unilocular; in Siphonodon with eight to twelve locules each locule secondarily transversely divided by secondary septa); dorsal bulge present in ovaries with apical septum. Gynophore sometimes present. Style usually single, simple (stylodia in Stackhousioideae sometimes two to five, free or connate at base), short (rarely long), hollow, or absent. Stigmas two to five, or one (in Pottingeria trilobate, with long spreading lobes), capitate or lobate, not or only little widened, commissural, non-papillate, Dry type. Male flowers with or without pistillodium.

Ovules Placentation usually axile to apical (rarely parietal or basal; in Pottingeria intrusively parietal). Ovules one to twelve (to numerous; in Pottingeria numerous) per carpel, usually anatropous (rarely amphitropous), ascending to pendulous (in Pottingeria sometimes horizontal), usually apotropous to pleurotropous (in Tripterygium epitropous), bitegmic, usually crassinucellar (rarely tenuinucellar). Micropyle usually bistomal (in Empleuridium endostomal; in Stackhousia exostomal; in Plagiopteron Z-shaped, zig-zag). Outer integument three to eight cell layers thick. Inner integument two to four cell layers thick. Megagametophyte monosporous, Polygonum type. Synergids sometimes with a filiform apparatus. Antipodal cells usually ephemeral (in Plagiopteron and Stackhousia proliferating to six to c. 20 cells). Endosperm development usually nuclear (in Pottingeria cellular). Endosperm haustoria absent. Embryogenesis caryophyllad or solanad (rarely asterad). Polyembryony (adventitious embryony) common in some species (embryos developing from inner integument).

Fruit Usually a loculicidal capsule, berry, drupe or samara (in Pottingeria a septicidal capsule with persistent stamens and placental vascular bundles; in Canotia loculicidal-septicidal; in Plagiopteron finally septicidal; in Pleurostylia a nut; in Stackhousioideae a schizocarp with two to five nutlike mericarps).

Seeds Seeds smooth or furrowed, with or without funicular aril or arilloid structure from hilum or exostoma (sometimes winged; wings in at least some species possibly homologous to aril; aril absent in Pottingeria). Seed coat usually exotegmic (occasionally reduced). Testa multiplicative (to 16 layers), usually lignified (sometimes vascularized). Exotesta with thick cuticle, often palisade (rarely tanniniferous and with thickened outer wall). Mesotestal cells sclerotic (rarely absent; in Pottingeria fibrous). Endotesta unspecialized? Exotegmen usually fibrous (consisting of tall lignified fibres). Endotegmen rarely persistent, tanniniferous. Perisperm not developed. Endosperm copious to sparse (in Brexia thin; in Pottingeria copious), oily, or absent. Embryo straight, well differentiated, usually with chlorophyll. Cotyledons two, large, sometimes connate. Germination phanerocotylar or cryptocotylar.

Cytology n = 8–10, 12, 14–17, 20, 23 (in Brexia n = 30, 32) – Polyploidy frequently occurring.

DNA Plastid gene infA present. Mitochondrial coxI intron present in Brexia.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), cyanidin, Ouratea catechins, diterpene and triterpene derivatives (quinone methides, celastrol, pristemerin, tingenin, etc.), gutta (trans-1,4-polyisoprene), tannins, proanthocyanidins (prodelphinidins), benzylisoquinoline alkaloids, pyrrolizidine alkaloids as aliphatic monocarboxylic esters, monoamine alkaloids (narcotic L(S)-(-)-alpha-aminopropiophenone, e.g. cathinone, cathine), sesquiterpene alkaloids (mayteine and 7-epi-mayteine, euonymine and 7-epi-euonymine, mekongensines, peritassines, etc., possessing cancer-inhibiting and other effects), saponins, d-norpseudoephedrin (cathin, in Catha edulis), maytansine (ansamycin macrolide; maytansinoids probably produced by endophytic actinobacteria), hexitols (dulcitol etc.), lupane lactones, and salacinol present. Ellagic acid and cyanogenic compounds not found. Nickel accumulated in species of Stackhousia.

Use Ornamental plants, medicinal plants, narcotics (Catha edulis), seed oils, fruits and seeds for food, insecticides, arrow poison, timber, carpentry, basketry.

Systematics Celastraceae are sister to Parnassiaceae.

Mortonia and Pottingeria (Pottingerioideae) form a sister-group to the remaining Celastraceae (Zhang & Simmons 2006). A provisional topology of Celastraceae is the following: [Pottingerioideae+[Microtropis clade+[Monimopetalum+[Stackhousioideae+[[Schaefferia clade+Celastroideae]+[American ‘Maytenus‘ clade+[[Gymnosporia clade+Old World ‘Maytenus‘ clade]+[Salaciopsis+[Brexioideae+[Hippocrateoideae+[Sarawakodendroideae+Salacioideae]]]]]]]]]]] (Bacon & al. 2016).

Nicobariodendron has dioecy, distichous leaves, no stipules, mucilage cells, racemose inflorescence, intrastaminal disc, two stamens, basal placentation and drupe (Vasudeva Rao & Chakrabarty 1985). It is known only from the type material from the Nicobar Islands.

Pottingerioideae Airy Shaw in Kew Bull. 28: 100. 1973

2/12. Pottingeria (1; P. acuminata; Naga Hills in Assam, northern Burma, northwestern Thailand), Mortonia (11; southwestern United States, northern Mexico). – Southeast Asia, southwestern United States, northern Mexico. Description of Pottingeria: Vessel elements in young stem with scalariform perforation plates. Leaves spiral. Stipules minute, cauline. Venation palmate. Leaf margin entire. Inflorescence fasciculate. Sepals and petals inserted abaxial on disc, free. Pistil composed of three connate carpels. Placentation intrusively parietal. Ovules numerous per carpel. Fruit a septicidal capsule with persistent stamens. Mesotesta fibrous. Endosperm copious. – Pottingeria was resolved as sister-group of Parnassiaceae on the rDNA tree (jacknife support 77%), but sister to Mortonia on the cpDNA tree (jacknife support 80%) of Zhang & Simmons (2006).

[Microtropis clade+[Monimopetalum+[Stackhousioideae+[[Schaefferia clade+Celastroideae]+[American ‘Maytenus‘ clade+[[Gymnosporia clade+Old World ‘Maytenus‘ clade]+[Salaciopsis+[Brexioideae+[Hippocrateoideae+[Sarawakodendroideae+Salacioideae]]]]]]]]]]

Microtropis clade

3/94. Zinowiewia (17; Mexico, Central America, Colombia, Venezuela), Microtropis (66; tropical Asia), Quetzalia (11; southern Mexico, Central America). – Tropical Asia, Mexico to Venezuela. – This clade was sister to the remaining Celastraceae in the combined analysis by Simmons, McKenna & al. (2012; see also Simmons, Bacon & al. 2012 and Simmons & Cappa 2013, etc.).

[Monimopetalum+[Stackhousioideae+[[Schaefferia clade+Celastroideae]+[American ‘Maytenus‘ clade+[[Gymnosporia clade+Old World ‘Maytenus‘ clade]+[Salaciopsis+[Brexioideae+[Hippocrateoideae+[Sarawakodendroideae+Salacioideae]]]]]]]]]

Monimopetalum clade

1/1. Monimopetalum (1; M. chinense; China). – Scandent shrub. Hairs absent. Leaves alternate. Leaf margin entire. Stipules pairwise, persistent. Inflorescence axillary, cymose. Flowers tetramerous. Disc globose, compressed. Stamens inserted on disc. Ovary quadrilocular. Ovules two per carpel, erect. Fruit a deeply quadrilobate loculicidal capsule with persistent wing-like petals. Aril small. – Monimopetalum chinense was sister to the remaining Celastraceae, according to Simmons & Cappa (2013).

[Stackhousioideae+[[Schaefferia clade+Celastroideae]+[American ‘Maytenus‘ clade+[[Gymnosporia clade+Old World ‘Maytenus‘ clade]+[Salaciopsis+[Brexioideae+[Hippocrateoideae+[Sarawakodendroideae+Salacioideae]]]]]]]]

Stackhousioideae Burnett, Outlines Bot.: 618, 1092, 1140. Feb 1835 [‘Stackhousidae’]

18/89. Wilczekra (1; W. congolensis; Central Africa); Hexaspora (1; H. pubescens; northeastern Queensland); Menepetalum (4; M. cassinoides, M. cathioides, M. salicifolium, M. schlechteri; New Caledonia), Dinghoua (1; D. globularis; northeastern Queensland), Apatophyllum (5; A. constablei, A. flavovirens, A. macgillivrayi, A. olsenii, A. teretifolium; southeastern Queensland, eastern New South Wales), Psammomoya (4; P. choretroides, P. ephedroides, P. grandiflora, P. implexa; southwestern Western Australia); Tripterococcus (4; T. brachystigma, T. brunonis, T. simplex, T. spathulatus; southwestern Western Australia), Macgregoria (1; M. racemigera; central Australia), Stackhousia (16; Malesia to New Guinea, Australia, Tasmania, New Zealand, Micronesia); Denhamia (10; northern and eastern Australia), Brassiantha (1; B. pentamera; New Guinea), Hedraianthera (1; H. porphyropetala; eastern Queensland, northeastern New South Wales), Dicarpellum (4; D. baillonianum, D. pancheri, D. paucisepalum, D. pronyense; New Caledonia), Hypsophila (3; H. dielsiana, H. halleyana, H. oppositiflora; northeastern Queensland). – Siphonodon (6; S. annamensis, S. australis, S. celastrineus, S. membranaceus, S. peltatus, S. pendulum; tropical Asia to New Guinea, eastern Queensland, northeastern New South Wales), Peripterygia (1; P. marginata; New Caledonia). Crossopetalum (c 25; tropical America), Xenodrys (1; X. micranthum; Madagascar). – Madagascar, Malesia to New Guinea, Australia, Tasmania, New Caledonia, New Zealand, Micronesia, tropical America. – The main part of Stackhousioideae s.lat. comprises “The Austral-Pacific clade”, according to Simmons & al. (2012) and Bacon & al. (2016). Hexaspora is sister to the remaining Stackhousioideae in Simmons & Cappa (2013) and the clade [[Menepetalum+Dinghoua]+[Apatophyllum+Psammomoya]] is successive sister-group to the remaining Stackhousioideae. [Menepetalum+[Apatophyllum+Psammomoya]] appears as a clade in Coughenour & al. (2010). [Macgregoria+Stackhousia] is sister to Tripterococcus. Brassiantha, Dicarpellum and Hypsophila form a Melanesian-NE Australian clade, perhaps together with Hedraianthera. [Peripterygia+Siphonodon] is sister-group to [Crossopetalum+Xenodrys] (Bacon & al. 2016).

[[Schaefferia clade+Celastroideae]+[American ‘Maytenus‘ clade+[[Gymnosporia clade+Old World ‘Maytenus‘ clade]+[Salaciopsis+[Brexioideae+[Hippocrateoideae+[Sarawakodendroideae+Salacioideae]]]]]]]

[Schaefferia clade+Celastroideae]

Schaefferia clade

5/32. Haydenoxylon (3; H. gentryi, H. haberianum, H. urbanianum; tropical South America), Gyminda (4; G. fimbrillata, G. latifolia, G. orbicularis, G. tonduzii; southern Mexico, Central America, the West Indies), Orthosphenia (1; O. mexicana; Mexico), Rzedowskia (1; R. tolantonguensis; Mexico), Schaefferia (23; tropical America). – Mexico, tropical America.

Celastroideae Burnett, Outlines Bot.: 621, 1140. Feb 1835 [‘Celastridae’] (under construction)

10/c 200. Celastrus (31; warm-temperate to tropical regions on both hemispheres), Tripterygium (1; T. wilfordii; eastern China, Taiwan); Paxistima (2; P. canbyi, P. myrsinites; North America), Wimmeria (12; Central America); Acanthothamnus (1; A. aphyllus; Mexico), Canotia (2; C. holacantha, C. wendtii; southwestern United States); Euonymus (c 130; temperate regions on the Northern Hemisphere, eastern Australia, Tasmania; non-monophyletic; incl. Glyptopetalum?, Torralbasia?, Xylonymus?), Glyptopetalum (c 20; tropical Asia), Torralbasia (1; T. cuneifolia; the West Indies), Xylonymus (1; X. versteeghii; western New Guinea).

[American ‘Maytenus‘ clade+[[Gymnosporia clade+Old World ‘Maytenus‘ clade]+[Salaciopsis+[Brexioideae+[Hippocrateoideae+[Sarawakodendroideae+Salacioideae]]]]]]

American Maytenus clade

5/145–160. Maytenus (15–24; Costa Rica, Panama, tropical and subtropical South America), Tricerma (6; T. obovatum, T. octogonum, T. orbiculare, T. texanum, T. viscifolium, T. vitis-idaeum; Florida, Texas, Mexico, Central America, the West Indies, tropical South America), Fraunhofera (1; F. multiflora; northeastern Brazil), Plenckia (3; P. bahiensis, P. microcarpa, P. populnea; Brazil, Paraguay, Bolivia, northwestern Argentina), Monteverdia (120–125; southern United States, Mexico, Central America, the West Indies, tropical South America). – Tropical and subtropical America.

[[Gymnosporia clade+Old World ‘Maytenus‘ clade]+[Salaciopsis+[Brexioideae+[Hippocrateoideae+[Sarawakodendroideae+Salacioideae]]]]]

[Gymnosporia clade+Old World ‘Maytenus‘ clade]

Gymnosporia clade

8?/>113? ‘Gymnosporia’ (>100; southern Spain, Madeira, the Canary Islands, northern to southern Africa, Madagascar, southwestern India, Sri Lanka, Malesia to eastern Queensland and islands in the Pacific; diphyletic), Lydenburgia (2; L. abbottii, L. cassinoides; Eastern Cape, Northern Province), Platypterocarpus (1; P. tanganyikensis; western Usambara Mountains in Uganda; possibly extinct), Allocassine (1; A. laurifolia; eastern Zimbabwe, southern Mozambique, Eastern Cape, KwaZulu-Natal, Swaziland), Cassine (3?; South Africa, Swaziland), Lauridia (2; L. reticulata, L. tetragona; South Africa, Swaziland), Maurocenia (1; M. frangula; Western Cape), Catha (3; C. abbottii, C. edulis, C. transvaalensis; eastern tropical and southern Africa, Madagascar, Yemen). – Tropical and subtropical regions in the Old World, tropical Asia to tropical Australia and New Caledonia, the West Indies, with their highest diversity on Madagascar.

Old World 'Maytenus' clade

6?/? Empleuridium (1; E. juniperinum; Caledon District in Western Cape), Pterocelastrus (4; P. echinatus, P. galpinii, P. rostratus, P. tricuspidatus; southeastern Africa, Zimbabwe, Mozambique, Malawi), ‘Maytenus’ pro parte (?; tropical and subtropical regions in the Old World), Mystroxylon (1; M. aethiopicum; eastern and southern Africa, Madagascar, the Mascarene Islands), Pseudosalacia (1; P. streyi; southern KwaZulu-Natal), Robsonodendron (2; R. eucleiforme, R. maritimum; South Africa, Swaziland).

[Salaciopsis+[Brexioideae+[Hippocrateoideae+[Sarawakodendroideae+Salacioideae]]]]

Salaciopsis

Salaciopsis (6; S. glomerata, S. longistyla, S. neocaledonica, S. sparsiflora, S. tapeinospermophylla, S. tiwakae; New Caledonia).

[Brexioideae+[Hippocrateoideae+[Sarawakodendroideae+Salacioideae]]]

Brexioideae Burnett, Outl. Bot.: 896, 1093, 1127. Jun 1835

15/108–123. Brexia (1 or 12; coastal regions in tropical East Africa, Madagascar, the Seychelles), ‘Polycardia’ (4; P. aquifolium, P. lateralis, P. libera, P. phyllanthoides; Madagascar; paraphyletic); Kokoona (10; Sri Lanka to Malesia), Lophopetalum (18–20; tropical Asia to tropical Australia); ‘Brexiella’ (c 10; Madagascar; non-monophyletic), ‘Euonymopsis’ (8; Madagascar; non-monophyletic), Elaeodendron (c 40; tropical and subtropical regions in the Old World, the West Indies), Pseudocatha (1; P. mandenensis; Madagascar), Halleriopsis (1; H. cathoides; Madagascar), Erythrostegia (2; E. grandis, E. integrifolia; Madagascar), Astrocassine (2–4; A. glomerata, A. pleurostylioides; Madagascar), Salvadoropsis (4; S. arenicola, S. grevicola, S. ludiifolia, S. paniculata; Madagascar), Pleurostylia (5; P. africana, P. capensis, P. opposita, P. pachyphlaea, P. serrulata; tropical and southern Africa, Madagascar, the Mascarene Islands, tropical Asia to New Guinea, northern Australia, New Caledonia), Hartogiopsis (1–2; H. trilobocarpa; Madagascar), Macrynodrys (1; M. mcphersonii; Madagascar).

[Hippocrateoideae+[Sarawakodendroideae+Salacioideae]]

Hippocrateoideae Lindl., Intr. Nat. Syst. Bot.: 120. 27 Sep 1830 [‘Hippocrateae’]

18/106. Plagiopteron (1; P. suaveolens; southern China, southern Burma, Thailand); Helictonema (1; H. velutinum; tropical Africa); Pristimera (21; tropical regions on both hemispheres); Cuervea (7; C. crenulata, C. hawkesii, C. integrifolia, C. isangiensis, C. jamaicensis, C. kappleriana, C. macrophylla; tropical West Africa, tropical America), Anthodon (4; A. decussatum, A. panamense, A. selloanum, A. trinerve; Central America, tropical South America), Arnicratea (3; A. cambodiana, A. ferruginea, A. grahamii; India to Southeast Asia), Bequaertia (1; B. mucronata; tropical Africa), Elachyptera (8; tropical Africa, Madagascar, tropical America), Apodostigma (1; A. pallens; tropical Africa, Madagascar), Reissantia (7; R. angustipetala, R. arborea, R. buchananii, R. cassinoides, R. indica, R. parviflora, R. setulosa; tropical regions in the Old World), Hylenaea (3; H. comosa, H. praecelsa, H. unguiculata; Central America, tropical South America), Semialarium (2; S. mexicanum, S. paniculatum; southern Mexico, Central America, tropical South America), Loeseneriella (16; tropical regions in the Old World), Hippocratea (16; tropical Africa, tropical America), Campylostemon (10; tropical Africa), Tristemonanthus (2; T. mildbraedianus, T. nigrisilvae; tropical West and Central Africa), Simicratea (1; S. welwitschii; Angola), Trochantha (2; T. graciliflora, T. preussii; tropical Africa). – Tropical regions on both hemispheres, with their highest diversity in tropical Africa. Flowers bisexual, pentamerous. Nectariferous disc extrastaminal. Stamens usually three. Anthers transversely dehiscent. Fruit a transversely flattened, deeply trilobate capsule. Testa with basal membranous wings or narrow stipes. – Plagiopteron was sister to the remaining Hippocrateoideae in the combined analysis by Simmons & al. (2001). Helictonema is sister to the remaining Hippocrateoideae “above” Plagiopteron (Coughenour 2010, 2011).

[Sarawakodendroideae+Salacioideae]

Sarawakodendroideae Savinov et Melikyan in Bot. Žurn. (Moscow and Leningrad) 87(7): 109. 29 Jul 2002

1/1. Sarawakodendron (1; S. filamentosum; Borneo). – Sarawakodendron is sister to Salacioideae, according to Coughenour & al. (2010). Median sepal abaxial. Two filaments shorter than remainder. Aril filaments supposedly homologous to spiral filaments in mucilaginous pulp in Salacioideae.

Salacioideae N. Hallé ex Thorne et Reveal in Bot. Rev. (Lancaster) 73: 181. 29 Jun 2007

5/c 270. Cheiloclinium (15; tropical America), Peritassa (c 20; Tobago, tropical South America), ‘Salacia’ (c 200; tropical and subtropical regions on both hemispheres; non-monophyletic; incl. Tontelea?), Tontelea (c 30; tropical America; in Salacia?), Salacighia (2; S. letestuana, S. linderi; tropical West and Central Africa to Angola), Thyrsosalacia (2; T. nematobrachion, T. viciflora; Central Africa). – Pantropical. Bisexual. Flowers pentamerous. Nectariferous disc extrastaminal. Stamens usually three. Anthers transversely dehiscent. Fruit a berry with arillate mucilaginous pulp.

Unplaced Celastraceae

Goniodiscus (1; G. elaeospermus; Brazil), Katafa (1; K. crassisepalum; Madagascar; in Brexioideae?), Ptelidium (2; P. ovatum, P. scandens; Madagascar; in Brexioideae?), Tetrasiphon (1; T. jamaicensis; Jamaica).

Phylogeny (simplified) of Celastraceae based on DNA sequence data and morphology (Simmons & al. 2001; Zhang & Simmons 2006).

Vegetative anatomy Phellogen? Vessel elements with simple perforation plates; lateral pits alternate or scalariform, simple or bordered pits. Vestured pits present in Ruptiliocarpon. Imperforate tracheary xylem elements thin-walled fibre tracheids? (tracheids absent) with simple (or bordered?) pits, non-septate. Wood rays uniseriate, homocellular. Axial parenchyma apotracheal, diffuse or diffuse-in-aggregates (or paratracheal?). Wood fluorescent. Sieve tube plastids Ss type. Nodes in Lepidobotrys 2:2, bilacunar with two leaf traces. Prismatic calciumoxalate crystals abundant. Druses absent (at least in flowers).

Leaves Alternate (distichous), basically palmately compound with one entire leaflet (unifoliolate), with ? ptyxis. Rhachis usually with long adaxial caducous stipulule (stipel; in Lepidobotrys two-veined) and articulated petiol(ul)e. Stipules usually caducous (rarely fused with petiole); leaf sheath absent. Petiole vascular bundle transection? Venation pinnate. Stomata laterocytic or paracytic. Cuticular wax crystalloids? Cristarque cells enveloped by vascular bundles. Leaf margin entire.

Inflorescence Terminal (often appearing leaf-opposite), raceme-like or spicate (sometimes congested).

Flowers Actinomorphic, small. Hypogyny. Sepals five, with imbricate quincuncial aestivation, in female flowers persistent, usually slightly connate at base. Petals five, with imbricate quincuncial aestivation, free. Nectariferous disc intrastaminal, in Lepidobotrys cupular (from receptacle), in Ruptiliocarpon tubular (from basally connate stamens).

Androecium Stamens 5+5, antepetalous longer than antesepalous. Filaments free (Lepidobotrys) or connate at base into tube (Ruptiliocarpon), free from tepals. Anthers basifixed to dorsifixed, versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Female flowers with 5+5 staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolporoidate (Lepidobotrys), shed as monads, ?-cellular at dispersal. Exine in Lepidobotrys crassitegillate-intectate, with intraluminar columellae; exine in Ruptiliocarpon tectate, with columellate infratectum, verrucate to fossulate/foveolate.

Gynoecium Pistil composed of two or three connate carpels. Ovary superior, bilocular or trilocular. Stylodia two or three more or less free and stigmas capitate, or style single simple short and stigma bilobate or trilobate, stigma type? Male flowers with pistillodium.

Ovules Placentation apical-axile. Ovules two per carpel, pachychalazal, collateral, anatropous, pendulous, epitropous, bitegmic, tenuinucellar. Micropyle bistomal. Outer integument seven to ten cell layers thick. Inner integument approx. four cell layers thick (Ruptiliocarpon). Integument multiplicative. Funicular obturator present above micropyle. Parietal tissue approx. ten cell layers thick. Megagametophyte monosporous, 8-nucleate (Polygonum-type?). Endosperm development? Endosperm haustoria? Embryogenesis?

Fruit A usually single-seeded septicidal capsule with mesocarp/exocarp separating from endocarp. Mesocarp with radially arranged scalariform fibres. Endocarp corneous, distinctly delimited. Columella persistent.

Seeds Carunculus inserted at apex of ovary locule. Seed coat exotegmic. Testa and tegmen multiplicative. Exotegmic cells in Ruptiliocarpon fibrous (not in Lepidobotrys), heavily thickened; remaining cells more or less crushed (in Lepidobotrys?). Endotegmen? Perisperm not developed. Endosperm absent. Embryo straight, oily, well differentiated, chlorophyll? Cotyledons two. Germination?

Parnassiales J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 110. 1846 [’Parnassieae’]; Lepuropetalaceae (Engl.) Nakai, Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov.]: 243. 20 Jul 1943

Distribution Temperate regions on the Northern Hemisphere, southeastern United States, Mexico, Uruguay, central Chile.

Habit Bisexual, annual succulent (Lepuropetalon) or perennial (Parnassia) herbs.

Vegetative anatomy Rhizome in Parnassia with cortex and medulla consisting of starch-filled parenchymatic cells. Phellogen? Young stems with separate vascular bundles. Pericycle six-layered, fibrous (Parnassia), or one- or two-layered, sclerenchymatic (Lepuropetalon). Secondary lateral growth absent? Vessel elements with simple perforation plates; lateral pits? Imperforate tracheary xylem elements? Wood rays? Axial parenchyma? Sieve tube plastids S type? Nodes 1:1, unilacunar with one leaf trace. Crystals? Druses absent (at least in flowers).

Leaves Alternate (spiral), simple, entire, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate or annular. Venation palmate or almost palmate, campylodromous (Parnassia) or acrodromous (Lepuropetalon), with veins proceeding to congested caducous foliar teeth. Stomata anomocytic. Cuticular wax crystalloids? Leaves without crystals. Young leaves in Parnassia with mucilaginous fringed appendages. Epidermis with distinct tanniniferous secretory cells (Lepuropetalon). Leaf margin serrate or entire.

Inflorescence Terminal, monochasium, or flowers solitary (basically reduced cyme).

Flowers Actinomorphic (Lepuropetalon) or weakly zygomorphic (Parnassia). Hypogyny (Parnassia) or half epigyny (Lepuropetalon). Floral receptacle distinctly hollow. Sepals (four or) five (to seven), with imbricate quincuncial aestivation, persistent, connate at base. Petals in Parnassia (four or) five (to seven), with imbricate quincuncial aestivation, persistent, free, inLepuropetalon rudimentary or absent. Nectaries intrastaminal, inserted at staminodial bases. Disc absent.

Androecium Stamens (four or) five (to seven), obdiplostemonous, antesepalous, alternipetalous. Filaments free from each other and from tepals. Anthers basifixed to ventrifixed, often versatile, usually tetrasporangiate (in Lepuropetalon disporangiate), extrorse (introrse as unripe and bending inwards one after the other during maturation), longicidal (dehiscent by longitudinal slits), often glandular, lobate (with apical glands on lobes). Tapetum secretory, with uninucleate or binucleate cells. Staminodia five, alternisepalous, antepetalous, intrastaminal trilobate to 15-lobate usually with apical glands on lobes (Parnassia) or extrastaminal simple (Lepuropetalon).

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolporate (rarely tetracolporate or syncolpate), shed as monads, bicellular at dispersal. Exine semitectate, with columellate? infratectum, reticulate.

Gynoecium Pistil composed of three or four (or five) connate antepetalous carpels, syncarpous at base and paracarpous above base; odd carpel abaxial. Ovary in Lepuropetalon semi-inferior, unilocular, in Parnassia superior, unilocular in upper part and multilocular in lower part. Style single, simple, very short or absent (Parnassia), or stylodia three or four (or five), very short, free (Lepuropetalon). Stigmas commissural (in Lepuropetalon capitate), papillate, Dry type. Pistillodium absent.

Ovules Placentation parietal (Lepuropetalon) or axile with T-shaped placentae (Parnassia). Ovules c. 25 to more than 100 per ovary, anatropous, horizontal, bitegmic (Parnassia) or unitegmic (Lepuropetalon), tenuinucellar. Micropyle bistomal (Parnassia). Outer integument two or three cell layers thick (Parnassia). Inner integument three or four cell layers thick. Integument in Lepuropetalon two cell layers thick. Parietal tissue one cell layer thick or absent. Nucellar cap absent? Megagametophyte monosporous, Polygonum type. Synergids with a filiform apparatus. Antipodal cells three, proliferating to approx. five cells or non-proliferating. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis asterad.

Seeds Aril absent. Testa in Parnassia winged. Raphe absent. Exotestal cells in Parnassia with thickened anticlinal walls. Endotestal cells in Lepuropetalon with heavily thickened walls. Tegmen multiplicative. Exotegmic cells tanniniferous, in Parnassia with U-shaped wall thickenings. Air spaces present in Parnassia between testa and embryo. Perisperm not developed. Endosperm one-layered or absent. Embryo straight, chlorophyll? Cotyledons two, short. Germination phanerocotylar?

Cytology n = 8, 9, 18 (Parnassia; Parnassia palustris: 2n = 17, 18, 27, 36, 32–37, 43–45 och 54); n = 8, 9, 23 (Lepuropetalon) – Allopolyploidy and autopolyploidy occurring in Parnassia.

Phytochemistry Insufficiently known. Flavonols and hexitols (dulcitol etc.) present. Ellagic acid and cyanogenic compounds not found.

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”The COM clade”

CELASTRALES Link

CELASTRACEAE R. Br.

LEPIDOBOTRYACEAE J. Léonard

PARNASSIACEAE Martinov