[Rubiales+[Plantaginales+Solanales]]


RUBIALES Juss. ex Bercht. et J. Presl

Berchtold et Presl, Přir. Rostlin: 256. Jan-Apr 1820 [‘Rubiaceae’]

Gentianales Juss. ex Bercht. et J. Presl, Přir. Rostlin: 248. Jan-Apr 1820 [‘Gentianeae’]; Gentiananae Thorne ex Reveal in Novon 2: 236. 13 Oct 1992

Habit Usually bisexual (rarely monoecious, gynomonoecious, polygamomonoecious, dioecious, gynodioecious, or functionally dioecious), evergreen trees, shrubs, suffrutices or lianas, perennial, biennial or annual herbs. Numerous species are xerophytes. Many are stem succulents. Some species are myrmecotrophic. Few species are epiphytic.

Vegetative anatomy Phellogen ab initio usually superficial (sometimes deeply seated). Secondary lateral growth normal or anomalous (from cylindrical cambium or concentric cambia). Endodermis often prominent. Vessel elements usually with simple (sometimes scalariform, rarely reticulate) perforation plates; lateral pits alternate, simple or bordered pits. Vestured pits present. Imperforate tracheary elements tracheids, fibre tracheids or libriform fibres with simple and/or bordered pits, usually non-septate (sometimes septate; also vasicentric tracheids). Wood rays uniseriate or multiseriate, homocellular or heterocellular, or absent. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty vasicentric, reticulate, scalariform, unilateral, confluent or banded (sometimes absent). Cambium and wood elements sometimes storied. Intraxylary phloem usually present. Sieve tube plastids S0 or Ss type. Nodes usually 1:1, unilacunar with on leaf trace, or 3:3, trilacunar with three traces (sometimes multilacunar, with several traces). Non-articulated branched or unbranched laticifers with white or bluish latex sometimes present. Prismatic calciumoxalate crystals as raphids, crystal sand, acicular crystals, styloids or druses often present.

Trichomes Hairs unicellular or multicellular, uniseriate or multiseriate, simple or branched, stellate, T-shaped, spiral, band-like, dendritic, candelabra-shaped, moniliform, papillose or lepidote, or absent; glandular hairs sometimes present.

Leaves Usually opposite (sometimes verticillate, rarely alternate), simple, usually entire (rarely lobed or modified into spines), sometimes coriaceous (rarely scale-like), usually with conduplicate or flat ptyxis. Stipules interpetiolar, intrapetiolar, sheathing or absent; leaves sometimes united simply by line across stem/branch; leaf sheath absent. Colleters consisting of secretory palisade layer surrounding elongate central axis; usually present on adaxial side of stipules, in leaf axils, on nodes and/or petiole bases, also on leaves, calyx and corolla. Petiole vascular bundle transection arcuate or annular. Venation usually pinnate (sometimes palmate or leaves uninerved). Stomata anomocytic, paracytic, anisocytic, or cyclocytic (sometimes parallelocytic, rarely actinocytic). Cuticular wax crystalloids? Domatia as pits, pockets or hair tufts (acarodomatia or bacteriodomatia rarely present), or absent. Epidermis with or without crystal idioblasts. Secretory cavities (laticifers) sometimes with latex. Epidermis and/or mesophyll with or without mucilage cells. Mesophyll with or without sclerenchymatous idioblasts. Leaf margin usually entire (rarely serrate or crenate). Extrafloral nectaries present in some species on adaxial surface of lamina.

Inflorescence Terminal or axillary, panicle, thyrse, umbel-, spike- or head-like, dichasial, corymb, pseudoverticillate or fasciculate cymose, or flowers solitary terminal or axillary.

Flowers Usually actinomorphic (rarely zygomorphic). Epicalyx usually absent. Hypogyny, epigyny or partial epigyny. Sepals (three or) four or five (to 16), with imbricate, convolute or valvate (sometimes open or contorted) aestivation, often persistent (sometimes accrescent), usually more or less connate (rarely free), often with adaxial colleters at base. Petals (three or) four or five (to 16), usually with contorted, imbricate or valvate (rarely imbricate or cochleate) aestivation, more or less connate into hypocrateromorphous, infundibuliform, campanulate or tubular corolla, often with adaxial hairs. Nectaries usually present, sometimes on abaxial side of ovary, sometimes as nectariferous glands. Intrastaminal, annular disc often present.

Androecium Stamens (one, three or) four or five (to 25), haplostemonous, antesepalous, alternipetalous. Filaments free or connate into tube (rarely connate in fascicles), free from corolla tube or epipetalous (filaments sometimes with nectariferous appendages of various shape, forming parts of gynostegium). Anthers usually free (sometimes connate and/or adnate to style into gynostegium), basifixed or dorsifixed, versatile or non-versatile, usually tetrasporangiate (sometimes disporangiate), usually introrse (occasionally extrorse or latrorse), usually longicidal (dehiscing by longitudinal slits; sometimes poricidal, dehiscing by apical pores). Tapetum usually secretory (rarely amoeboid-periplasmodial). Staminodia usually absent (sometimes one to four, alternating with fertile stamens; female flowers often with staminodia). Pollinaria present in many Apocynaceae (especially in Asclepiadoideae), consisting of pollinia and translator.

Pollen grains Microsporogenesis usually simultaneous (sometimes successive). Pollen grains (2–)3(–6)-colporate or (1–)3(–6)-porate (rarely colpate, pororate, polyporate or syncolpate), usually shed as monads (sometimes tetrads, massulae or pollinia), bicellular or tricellular at dispersal. Exine tectate or semitectate (rarely intectate), usually with columellate (sometimes granular) infratectum, perforate, reticulate, microreticulate, striate, scabrate, psilate, fossulate, spinulate, verrucate, echinulate, granulate, or smooth.

Gynoecium Pistil composed of usually two (rarely three to eight or up to 16) connate carpels (carpels sometimes partially free, or secondarily free in ovary region). Ovary superior, inferior or semi-inferior, usually bilocular (sometimes unilocular, rarely trilocular to novemlocular, or divided by secondary septa). Style single, simple, or stylodia two (to five; sometimes connate above only), sometimes persistent (rarely absent). Stigma capitate, punctate, clavate, truncate, or bilobate (to quinquelobate; sometimes subpeltate, or swollen at apex and constricted in central part and in lower part ring-like), papillate or non-papillate, Dry or Wet type. Pistillodium usually absent (male flowers often with pistillodium).

Ovules Placentation axile or apical (when ovary bilocular or multilocular) or parietal (when ovary unilocular; sometimes intrusively parietal, rarely basal or free central). Ovules one to numerous per carpel, anatropous or amphitropous (rarely campylotropous, orthotropous, circinotropous or hypertropous), usually pendulous (sometimes horizontal or ascending), apotropous or epitropous, usually unitegmic (rarely ategmic), usually tenuinucellar (often reduced tenuinucellar; rarely crassinucellar or pseudocrassinucellar or megasporangium absent). Endothelium absent. Megagametophyte usually monosporous, Polygonum type (rarely disporous, Allium type, or tetrasporous, Drusa type). Synergids sometimes with a filiform apparatus. Antipodal cells sometimes persistent, occasionally proliferating and/or haustorial. Endosperm development usually ab initio nuclear (rarely cellular). Endosperm haustorium micropylar, chalazal (rarely lateral) or absent. Embryogenesis caryophyllad or solanad (rarely onagrad).

Fruit A loculicidal and/or septicidal (rarely denticidal) capsule, a berry, a drupe, or a schizocarp with usually two follicular, nutlike, drupaceous or baccate mericarps (rarely a follicle or a pyxidium, or a pseudofruit consisting of connate ovaries), calyx often persistent.

Seeds Aril sometimes present. Testa usually multiplicative (seed coat sometimes thin or absent). Exotestal cell walls usually thickened (theoidal exotestal thickenings). Mesotesta sometimes with flattened crystalliferous cells. Endotesta often crushed. Perisperm not developed. Endosperm copious to sparse, oily (sometimes with starch or hemicellulose), sometimes ruminate, or absent. Embryo large or small, straight or curved, well differentiated (rarely rudimentary, undifferentiated), with or without chlorophyll. Cotyledons two. Germination phanerocotylar or cryptocotylar.

Cytology x = (5–)8–15 – Protein crystalloids present in nucleus.

DNA Mitochondrial intron coxII.i3 present or absent.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin [rare]), flavones, O-methylated flavones, cyanidin, delphinidin, Route I iridoids (also secoiridoids), Route II decarboxylated iridoids, Group I carbocyclic iridoids (theviridoside, daphylloside, plumieride, geniposide, scandoside, monotropein, gardenoside), Group II carbocyclic iridoids (shantziside), Group VI secoiridoids (secologanin), Group VII secoiridoids (sweroside, swertiamarin, gentiopicroside), Group IX secoiridoids (indole alkaloids of corynanthe type, aspidosperma and iboga type indole alkaloids, ipecacalkaloids), Group X secoiridoids (desoxyloganin, loganin, ketologanin, antirrhide, gardoside, iridoid pyridine alkaloids), iridoid coumarins (asperuloside), cardenolides, triterpenes (in latex), tannins, ursolic acid, caffeic acid, pyrrolizidine alkaloids as aliphatic monocarboxylic esters and esters of arylic and aralkylic acids, camptothecine and other C17 indole alkaloids, isoquinoline alkaloids, steroid alkaloids, tryptophane-derived alkaloids, toxic (cardiotonic) steroid glycosides, cyanogenic compounds (rare), saponins, anthraquinones (usually shikimic acid derived), xanthones, and arbutin present. Ellagic acid not found.

Systematics Rubiales may be sister to [Lamiales+Solanales], although the support is low.

Rubiaceae are sister to the [Gentianaceae+[Loganiaceae+[Gelsemiaceae+Apocynaceae]]] clade. Potential synapomorphies of this clade are presence of intraxylary phloem; late corolla tube formation; postgenital syncarpy; and presence of Route I secoiridoids (Stevens 2001 onwards). Gelsemiaceae and Apocynaceae share the potential morphological synapomorphies nodes 1:1 and flattened seeds.

Cladogram of Rubiales based on DNA sequence data (Soltis & al. 2011; Frasier 2009; etc.).

APOCYNACEAE Juss.

( Back to Rubiales )

de Jussieu, Gen. Plant.: 143. 4 Aug 1789 [’Apocineae’], nom. cons.

Asclepiadaceae Borkh., Bot. Wörterb. 1: 37. 1797 [‘Asclepiadeae’], nom. cons.; Vincaceae Vest, Anleit. Stud. Bot.: 273, 299. 1818 [‘Vincoideae’]; Apocynales R. Br. ex Bercht. et J. Presl, Přir. Rostlin: 249. Jan-Apr 1820 [‘Apocineae’]; Asclepiadales R . Br. ex Bercht. et J. Presl, Přir. Rostlin: 249. Jan-Apr 1820 [‘Asclepiadeae’]; Cerberaceae Martinov, Tekhno-Bot. Slovar: 119. 3 Aug 1820 [‘Cerberides’]; Pacouriaceae Martinov, Tekhno-Bot. Slovar: 447. 3 Aug 1820 [‘Pacurides’]; Plumeriaceae Horan., Prim. Lin. Syst. Nat.: 70. 2 Nov 1834 [‘Plumeriaceae (Apoocyneae)’]; Stapeliaceae Horan., Prim. Lin. Syst. Nat.: 70. 2 Nov 1834 [‘Stapeliaceae (Asclepiadeae)’]; Ophioxylaceae Mart., Consp. Regn. Veg.: 24. Sep-Oct 1835 [‘Ophioxyleae’]; Cynanchaceae G. Mey., Chloris Han.: 245, 251. Jul-Aug 1836 [‘Cynancheae’]; Asclepiadopsida Brongn., Enum. Plant. Mus. Paris: xxiii, 51. 12 Aug 1843 [’Asclepiadineae’]; Vincales Horan., Char. Ess. Fam.: 111. 30 Jun 1847 [’Vincastra’]; Willughbeiaceae J. Agardh, Theoria Syst. Plant.: 256. Apr-Sep 1858 [‘Willughbejieae’]; Carissaceae Bertol. in Nuevo Giorn. Bot. Ital. 23: 212. 8 Jan 1891; Periplocaceae (Kostel.) Schltr. in K. Schum. et C. A. G. Lauterb., Nachtr. Fl. Schutzgeb. Südsee: 351. 16 Nov 1905, nom. cons.

Genera/species 340–350/4.350–4.500

Distribution Cosmopolitan except polar areas and cold-temperate regions.

Fossils Unknown.

Habit Usually bisexual (rarely functionally dioecious), evergreen trees, shrubs or lianas (sometimes perennial or annual herbs). Numerous species are xerophytic and many are stem succulents. Some species (e.g. Dischidia rafflesiana) are myrmecotrophic with ant symbiosis (ant colonies present in leaves).

Vegetative anatomy Endomycorrhiza Arum type, Paris type or intermediate types. Phellogen ab initio usually superficial (in, e.g., Rhazya deeply seated). Secondary lateral growth normal or anomalous (from a cylindrical cambium or concentric cambia). Cambium sometimes storied. Pericyclic fibres usually absent. Endodermis often prominent. Vessel elements with usually simple (sometimes scalariform) perforation plates; lateral pits alternate, bordered pits. Vestured pits present. Imperforate tracheary elements tracheids or fibre tracheids with simple or bordered pits, usually 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, or banded, or absent. Wood elements sometimes storied. Intraxylary phloem (diffuse) usually present. Sieve tube plastids S type. Nodes usually 1:1, unilacunar with one leaf trace (rarely 3:3, trilacunar with three traces). Articulated or non-articulated branched or unbranched laticifers with white or bluish latex frequent (absent in, e.g., Nerium). Prismatic calciumoxalate crystals abundant; crystal sand or druses present in some representatives.

Trichomes Hairs unicellular or multicellular, simple or branched, sometimes dendritic, or absent.

Leaves Usually opposite (sometimes verticillate, rarely alternate), simple, usually entire (rarely lobed or transformed into spines), usually with conduplicate or flat ptyxis. Stipules absent (small, interpetiolar or cauline stipule-like processes sometimes present); leaf sheath absent. Axillary colleters often present (in association with stipules). Petiole vascular bundle transection arcuate?; petiole bundles sometimes adaxial. Venation usually pinnate (sometimes palmate or leaves one-veined). Stomata anomocytic, paracytic, anisocytic, cyclocytic, or parallelocytic (rarely actinocytic). Cuticular wax often as crust. Domatia as pits, pockets or hair tufts, or absent. Epidermis with or without crystal idioblasts. Secretory cavities (laticifers) with latex. Mesophyll with or without sclerenchymatous idioblasts. Leaf margin usually entire (rarely serrate). Extrafloral nectaries present on petiole and adaxial surface of lamina in numerous species.

Inflorescence Terminal or axillary, umbel-like, panicle, dichasial, etc. cymose, or flowers solitary axillary.

Flowers Actinomorphic. Hypogyny or partial epigyny. Sepals usually five (sometimes four), with imbricate quincuncial or valvate aestivation, more or less connate, with adaxial colleters at base. Petals usually five (sometimes four), usually with sinistrally contorted (rarely imbricate or valvate) aestivation, more or less connate; upper part of corolla tube (above staminal insertion point) formed by postgenital fusion; corona often present as scale-like appendages in corolla tube mouth. Nectaries usually present, sometimes on abaxial side of ovary. Intrastaminal disc often present.

Androecium Stamens (four or) five, haplostemonous, antesepalous, alternipetalous. Filaments short, free or connate into tube, free from or adnate to corolla tube; filaments in Secamonoideae? and Asclepiadoideae usually with nectariferous appendages of various shape, forming parts of gynostegium. Anthers free, often connivent, or more or less connate and sometimes adnate to upper part of style and stigma into gynostegium, with or without apical, lateral or dorsal appendages, basifixed or dorsifixed, non-versatile, disporangiate (in Asclepiadoideae) or tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits) or poricidal (dehiscing by apical pores); anthers often sterile below and prolonged into acute tip; connective often prolonged at apex, in Asclepiadoideae (and Secamonoideae?) with wing-like and membranous appendages forming part of corona complex. Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous or successive. Pollen grains (2–)3–6-colporate or (2–)3–6-porate (sometimes with pseudocolpi), shed as monads, tetrads (tetrahedral, rhomboidal or linear), massulae or pollinia, bicellular or tricellular at dispersal (usually transported within a foam-like substance). Exine tectate, with granular or columellate infratectum, perforate or psilate, fossulate, verrucate, granulate, or smooth (occasionally microreticulate or scabrate).

Gynoecium Pistil composed of usually two (rarely three to eight) connate carpels, or carpels secondarily free in ovary region, connate usually only in stylar part (often exclusively in upper[most] part of style); connation of carpels congenital or postgenital; carpels sometimes collateral; carpels sometimes transversely orientated. Ovary superior or semi-inferior, usually bilocular (sometimes unilocular); locules usually free. Stylodia two, connate entirely or only in upper(most) part; stylar apex usually swollen, pentagonal, with cells secreting pollen-adhering viscid substance consisting of terpenoids and polysaccharides. Stigma usually with terminal stigmatic head, swollen at apex, constricted in central part and with receptive ring, hair ring or membrane in lower part; pollen-receptive surfaces lateral, alternating with stamens; stigmatic surfaces papillate or non-papillate, Dry or Wet type. Gynostegium often present, developed through postgenital fusion of anther connective and stigmatic head. Pistillodium absent. Secondary pollen presentation often occurring (pollen grains deposited on apex of stigmatic head).

Ovules Placentation axile (ventral in each locule) or apical (when ovary bilocular or multilocular) or parietal (when ovary unilocular). Ovules two to numerous (rarely one) per carpel, anatropous, amphitropous or hemitropous (rarely orthotropous), usually pendulous, unitegmic, usually tenuinucellar (reduced tenuinucellar, with meiocyte semi-inferior; sometimes pseudocrassinucellar). Integument six to nine cell layers thick. Endothelium absent (always?). Megagametophyte monosporous, Polygonum type. Synergids sometimes with a filiform apparatus. Antipodal cells sometimes persistent. Endosperm development ab initio nuclear. Endosperm haustorium micropylar?; chalazal haustorium usually absent. Embryogenesis caryophyllad or solanad.

Fruit A capsule, a berry, a drupe, or a schizocarp with two follicular, nutlike, drupaceous or baccate (often divergent) mericarps.

Seeds. Aril sometimes present. Testa winged or unwinged, usually multiplicative, often with terminal coma consisting of long silky hairs (seed coat sometimes absent). All exotestal cell walls usually thickened (in, e.g., Periploca non-thickened). Mesotesta sometimes with flattened crystalliferous cells. Endotesta? Perisperm not developed. Endosperm copious to sparse, oily, sometimes ruminate, or absent. Embryo straight, well differentiated, with or without chlorophyll. Cotyledons two, flat, folded or in-rolled. Germination phanerocotylar or cryptocotylar. Seedling leaves sometimes alternate.

Cytology x = (6–)9–11(–14) (23) – Polyploidy frequently occurring. Protein crystalloids present in nucleus.

DNA Mitochondrial intron coxII.i3 lost in Catharantus. Mitochondrial coxI intron usually present.

Phytochemistry Flavonols (kaempferol, quercetin), O-methylated flavones, cyanidin, delphinidin, Route I carbocyclic iridoids (theviridoside, daphylloside, plumieride), Route II decarboxylated iridoids, Group VI secoiridoids (secologanin), Group VII secoiridoids (sweroside), Group IX secoiridoids (indole alkaloids of corynanthe-, aspidosperma- and iboga-type indole alkaloids), Group X secoiridoids (desoxyloganin, loganin, ketologanin, iridoid pyridine alkaloids), cardiac glycosides (cardenolides), triterpenes (in latex), tannins, ursolic acid, caffeic acid, pyrrolizidine alkaloids as aliphatic monocarboxylic esters and esters of arylic and aralkylic acids, steroidal pregnane pseudoalkaloids, toxic (cardiotonic) steroidal glycosides, tryptophane-derived alkaloids, and saponins present. Cyanogenic compounds very rare. Ellagic acid not found. Aluminium accumulated in some species.

Use Ornamental plants, medicinal plants (Cataranthus roseus, Rauvolfia, etc.), arrow poison, perfumes (Plumeria etc.), timber, latex (gums from Carpodinus, Funtumia elastica, Hancornia speciosa, Landolphia, Mascarenhasia), fruits (Carissa carandas).

Systematics Apocynaceae are sister to Gelsemiaceae.

A plausible topology for the Apocynaceae phylogeny is the following (Livshultz & al. 2007; Lens & al. 2008).

[Aspidospermateae+[Alstonieae+[[Vinceae+[Willughbeeae+Tabernaemontaneae]]+[Diplorhynchus-Stephanostegia clade+Melodineae+Amsonia+[Condylocarpinae+[Chilocarpeae+[Alyxieae+[Plumerieae+[Carisseae+[Wrightieae+[Nerieae+[Malouetieae+[Rhabdadenia+[Periplocoideae+[Apocyneae+[New World clade+[Baisseeae+[Secamonoideae+Asclepiadoideae]]]]]]]]]]]]]]]]]]

Aspidospermateae Miers, Apocyn. S. Amer.: 7. Mai-Jun 1878 [‘Aspidospermeae’]

4/75–80. Aspidosperma (c 70; tropical America), Anechites (1; A. nerium; tropical America), Geissospermum (5; Brazil), Haplophyton (1; H. cimicidum; Mexico, Guatemala). – Tropical America. Uniseriate wood rays absent. Seed in Haplophyton with coma. – Aspidospermateae are sister-group to the remaining Apocynaceae.

[Alstonieae+[[Vinceae+[Willughbeeae+Tabernaemontaneae]]+[Diplorhynchus-Stephanostegia clade+Melodineae+Amsonia+[Condylocarpinae+[Chilocarpeae+[Alyxieae+[Plumerieae+[Carisseae+[Wrightieae+[Nerieae+[Malouetieae+[Rhabdadenia+[Periplocoideae+[Apocyneae+[New World clade+[Baisseeae+[Secamonoideae+Asclepiadoideae]]]]]]]]]]]]]]]]]

Alstonieae G. Don, Gen. Hist. 4: 70, 86. 1837-8 Apr 1838

5/50–55. Alstonia (40–45; tropical Africa, Madagascar, southern China, tropical Asia, Melanesia, islands in the western Pacific, one species, A. longifolia, in Central America), Laxoplumeria (3; eastern Peru, Brazil), Microplumeria (1; M. anomala; Amazonian Brazil), Strempeliopsis (2; Cuba, Jamaica), Dyera (2–3; West Malesia). – Nearly pantropical. Uniseriate wood rays absent.

[[Vinceae+[Willughbeeae+Tabernaemontaneae]]+[Diplorhynchus-Stephanostegia clade+Melodineae+Amsonia+[Condylocarpinae+[Chilocarpeae+[Alyxieae+[Plumerieae+[Carisseae+[Wrightieae+[Nerieae+[Malouetieae+[Rhabdadenia+[Periplocoideae+[Apocyneae+[New World clade+[Baisseeae+[Secamonoideae+Asclepiadoideae]]]]]]]]]]]]]]]]

[Vinceae+[Willughbeeae+Tabernaemontaneae]]

Vinceae Duby, Bot. Gall. 1: 324. 12-14 Apr 1828

7/145–150. Petchia (8; Cameroun, Madagascar, the Comoro Islands, Sri Lanka), Rauvolfia (c 80; tropical regions on both hemispheres), Catharanthus (8; Madagascar, India, Sri Lanka), Vinca (7; Europe, the Mediterranean, North Africa and eastwards to Central Asia), Ochrosia (40–43; the Mascarene Islands, the Seychelles, Southeast Asia, Malesia and eastwards to tropical Australia, islands in western Pacific), Kamettia (1; K. caryophyllata; southern India), Rhazya (1; R. stricta; southeastern Europe and eastwards to the Arabian Peninsula and Pakistan). – Tropical regions on both hemispheres. – Vinceae are sister to [Willughbeeae+Tabernaemontaneae]].

[Willughbeeae+Tabernaemontaneae]

Willughbeeae A. DC. in A. P. de Candolle et A. L. P. P. de Candolle, Prodr. 8: 318. Mar (med.) 1844 [‘Willughbeiae’]

18/c 145. Ancylobotrys (7; tropical and southern Africa, Madagascar), Bousigonia (2; Southeast Asia), Chamaeclitandra (1; C. henriquesiana; tropical Africa), Clitandra (1; C. cymulosa; tropical Africa), Couma (6; northeastern South America), Cyclocotyla (1; C. congolensis; Central Africa), Cylindropsis (1; C. parvifolia; tropical West and Central Africa), Dictyophleba (5; tropical Africa), Hancornia (4; Brazil), Lacmellea (c 20; tropical South America), Landolphia (c 55; tropical and southern Africa, the Mascarene Islands, tropical America), Leuconotis (5; Malesia), Orthopichonia (6; tropical West and Central Africa), Pacouria (2; tropical America), Parahancornia (6; tropical South America), Saba (3; tropical Africa, Madagascar), Vahadenia (2; tropical West and Central Africa), Willughbeia (16; tropical Asia). – Pantropical. – Willughbeeae may be sister to [Kopsia+Tabernaemontaneae].

Tabernaemontaneae G. Don, Gen. Hist. 4: 70, 87. 1837-8 Apr 1838

17/c 180. Kopsia (22; Southeast Asia, West Malesia, the Carolines); Molongum (3; tropical South America), Schizozygia (1; S. coffeoides; tropical East Africa, the Comoro Islands), Ambelania (3–4; tropical South America), Callichilia (7; tropical Africa), Tabernaemontana (c 110; tropical regions on both hemispheres), Calocrater (1; C. preussii; tropical West and Central Africa), Carvalhoa (1; C. campanulata; eastern and southeastern Africa), Crioceras (1; C. dipladeniiflorus; Gabon to Angola), Macoubea (3; tropical America), Mucoa (2; tropical South America), Neocouma (2; northern South America), Rhigospira (1; R. quadrangularis; northern South America, the Andes), Spongiosperma (6; northern South America), Tabernanthe (2; Central Africa), Voacanga (12; tropical and subtropical regions in the Old World), Woytkowskia (2; tropical South America). – Pantropical. Calyx with colleters at base. Filaments absent. Anthers connivent, with lignified basal appendages (nectar guides). Nectaries paired and fused with stigmatic head complex. Pollen grains sometimes porate. Ovary often apocarpous (carpels more or less free). Stigma usually with stigmatic head complex consisting of apical quinquelobate stigmatic crest and thickened basal flange. Placentation axile or parietal. Fruit a berry, a drupe or a follicle. Seed sometimes arillate, sometimes ruminate. x = (9) 10, 11 (23). Indole alkaloids often present. –Kopsia may be sister to the remaining Tabernaemontaneae.

[Diplorhynchus-Stephanostegia clade+Melodineae+Amsonia+Hunterieae+[Condylocarpinae+[Chilocarpeae+[Alyxieae+[Plumerieae+[Carisseae+[Wrightieae+[Nerieae+[Malouetieae+[Rhabdadenia+[Periplocoideae+[Apocyneae+[New World clade+[Baisseeae+[Secamonoideae+Asclepiadoideae]]]]]]]]]]]]]]]

Diplorhynchus-Stephanostegia clade

3/4. Diplorhynchus (1; D. condylocarpon; tropical and southern Africa), Pycnobotrya (1; P. nitida; Central Africa), Stephanostegia (2; Madagascar). – Tropical and southern Africa, Madagascar. The sister-group relationships of this clade are not resolved.

Melodineae G. Don, Gen. Hist. 4: 71, 101. 1837-8 Apr 1838

2/c 20. Craspidospermum (1; C. verticillatum; Madagascar), Melodinus (c 20; tropical Asia, Pacific islands). – Madagascar, tropical Asia, islands in the Pacific. The sister-group relationships of Melodineae are not resolved.

Amsonia clade

1/c 20. Amsonia (c 20; Japan, North America). – The sister-group relationships of Amsonia are not resolved.

Hunterieae Miers, Apocyn. S. Amer.: 6. Mai-Jun 1878

4/10. Gonioma (2; G. kamassi: Western and Eastern Cape, KwaZulu-Natal, Swaziland; G. malagasy: southwestern Madagascar), Hunteria (12; tropical Africa), Picralima (1; P. nitida; tropical West and Central Africa), Pleiocarpa (5; tropical Africa). – Tropical and southern Africa, Madagascar. – The sister-group relationships of Hunterieae are not resolved.

[Condylocarpinae+[Chilocarpeae+[Alyxieae+[Plumerieae+[Carisseae+[Wrightieae+[Nerieae+[Malouetieae+[Rhabdadenia+[Periplocoideae+[Apocyneae+[New World clade+[Baisseeae+[Secamonoideae+Asclepiadoideae]]]]]]]]]]]]]]

Condylocarpinae Pichon ex Leeuwenb. in Wageningen Agric. Univ. Pap. 94(3): 56. 19 Aug 1994

1/7. Condylocarpon (7; Nicaragua and southwards to Brazil).

[Chilocarpeae+[Alyxieae+[Plumerieae+[Carisseae+[Wrightieae+[Nerieae+[Malouetieae+[Rhabdadenia+[Periplocoideae+[Apocyneae+[New World clade+[Baisseeae+[Secamonoideae+Asclepiadoideae]]]]]]]]]]]]]

Chilocarpeae Pichon ex Leeuwenb. in Wageningen Agric. Univ. Pap. 94(3): 54. 19 Aug 1994

1/13. Chilocarpus (13; tropical Asia).

[Alyxieae+[Plumerieae+[Carisseae+[Wrightieae+[Nerieae+[Malouetieae+[Rhabdadenia+[Periplocoideae+[Apocyneae+[New World clade+[Baisseeae+[Secamonoideae+Asclepiadoideae]]]]]]]]]]]]

Alyxieae G. Don, Gen. Hist. 4: 71, 96. 1837-8 Apr 1838

4–5/110–115. Alyxia (105–110; tropical Asia, western Pacific islands, the Hawaiian islands; incl. Pteralyxia?), Pteralyxia (2; the Hawaiian Islands; in Alyxia?), Lepinia (4; New Guinea, the Solomon Islands, the Caroline Islands, the Marquesas, Tahiti, Moorea), Lepiniopsis (2; Central and East Malesia to New Guinea, Micronesia), Plectaneia (3; Madagascar). – Madagascar, tropical Asia to the Solomon Islands, Micronesia and Polynesia. Pollen grains irregularly-shaped, 2- or 3-porate. Ectoapertures with thickened margins.

[Plumerieae+[Carisseae+[Wrightieae+[Nerieae+[Malouetieae+[Rhabdadenia+[Periplocoideae+[Apocyneae+[New World clade+[Baisseeae+[Secamonoideae+Asclepiadoideae]]]]]]]]]]]

Plumerieae E. Mey., Comm. Plant. Afr. Austr. 2: 188. 1-8 Jan 1838

10/56–57. Allamanda (13–14; tropical America), Cameraria (2; the West Indies), Cerbera (2; coasts of the Indian and western Pacific Oceans), Cerberiopsis (2; New Caledonia), Himatanthus (13; South America), Mortoniella (1; M. pittieri; Central America), Plumeria (8; tropical America), Skytanthus (2; Brazil, Chile), Thevetia (8; tropical America), Vallesia (5; tropical America).

[Carisseae+[Wrightieae+[Nerieae+[Malouetieae+[Rhabdadenia+[Periplocoideae+[Apocyneae+[New World clade+[Baisseeae+[Secamonoideae+Asclepiadoideae]]]]]]]]]]

Carisseae Dumort., Anal. Fam. Plant.: 26. 1829

2/12. Acokanthera (5; eastern and southern Africa, Yemen), Carissa (7; tropical and southern Africa, Madagascar, the Mascarene Islands, tropical Asia to tropical Australia and New Caledonia). – Tropical regions in the Old World eastwards to New Caledonia.

[Wrightieae+[Nerieae+[Malouetieae+[Rhabdadenia+[Periplocoideae+[Apocyneae+[New World clade+[Baisseeae+[Secamonoideae+Asclepiadoideae]]]]]]]]]

Corolla usually dextrorsely contorted. Anthers with lignified basal appendages, adnate to stylar head. Retinaculum present (staminal portion consisting of trichomes and attaching to stylar head). Pollen grains porate. Gynoecium apocarpous. Stylar head radially differentiated. Fruit a follicle. Coma (hair tuft) usually present on chalazal end of testa. Iridoids absent. – Gynostegium formed by postgenital close connection between androecial and gynoecial development. Special pollination mechanism consisting of gynostegium in combination with lignified nectar guides.

Wrightieae G. Don, Gen. Hist. 4: 70, 85. 1837-8 Apr 1838

3/c 30. Wrightia (c 25; tropical and subtropical regions in the Old World; non-monophyletic?); Stephanostema (1; S. stenocarpum; Tanzania), Pleioceras (5; tropical Africa). – Tropical and subtropical regions in the Old World. Corolla sinistrorsely contorted.

[Nerieae+[Malouetieae+[Rhabdadenia+[Periplocoideae+[Apocyneae+[New World clade+[Baisseeae+[Secamonoideae+Asclepiadoideae]]]]]]]]

Nerieae Baill., Hist. Plant. 10: 166, 198. Sep-Oct 1889

5–6/c 70. Nerium (1; N. oleander; the Mediterranean, southwestern Asia; geographic origin unknown), Adenium (1 or 5; northeastern, tropical and southern Africa, southern Arabian Peninsula, Socotra); Strophanthus (38; tropical and southern Africa, Madagascar, tropical Asia), Isonema (3; tropical West and Central Africa), ‘Alafia’ (23; tropical Africa, Madagascar; paraphyletic), Farquharia (1; F. elliptica; southern Nigeria; in Alafia?). – The Mediterranean, tropical and southern Africa, Madagascar, the Arabian Peninsula, Socotra, southwestern and tropical Asia.

[Malouetieae+[Rhabdadenia+[Periplocoideae+[Apocyneae+[New World clade+[Baisseeae+[Secamonoideae+Asclepiadoideae]]]]]]]

Malouetieae Müll.-Arg. in C. F. P. von Martius, Fl. Bras. 6(1): 5, 89. 30 Jul 1866

11/c 90. Galactophora (6–7; tropical South America); Pachypodium (c 25; southern and southeastern Africa, Madagascar), Neobracea (4; Cuba, Bahamas), Spirolobium (1; S. cambodianum; Thailand, Indochina, the Malay Peninsula), Mascarenhasia (8; Madagascar, one species also in eastern and southern Africa), Holarrhena (4; tropical Africa, tropical Asia), Malouetia (c 20; tropical Africa, tropical America), Funtumia (2; tropical Africa), Kibatalia (c 15; Southeast Asia and eastwards to the Philippines), Allowoodsonia (1; A. whitmorei; the Solomon Islands), Carruthersia (3; the Philippines and eastwards to the Solomon Islands, Fiji to Tonga). – Pantropical. Micropylar coma often absent. – Galactophora is sister to the remaining Malouetieae.

[Rhabdadenia+[Periplocoideae+[Apocyneae+[New World clade+[Baisseeae+[Secamonoideae+Asclepiadoideae]]]]]]

Rhabdadenia clade

1/5. Rhabdadenia (5; the West Indies, tropical South America). – Vessels grouped in radial multiples. Tracheids absent. Wood fibres parenchymatous and very thin-walled. Retinaculum similar to that in Wrightieae, Nerieae and Malouetieae. – Rhabdadenia is recognized as basal to the Periplocoideae-Asclepiadoideae clade, according to analyses by Livshultz (2010), which is consistent with its morphology.

[Periplocoideae+[Apocyneae+[New World clade+[Baisseeae+[Secamonoideae+Asclepiadoideae]]]]]

Periplocoideae Endl., Gen. Plant.: 587. Aug 1838 [’Periploceae’]

37/c 170. Phyllanthera (2; tropical Asia and eastwards to islands in the Pacific); Atherandra (2; Southeast Asia, West Malesia), Atherolepis (3; Southeast Asia), Baroniella (8; Madagascar), Baseonema (1; B. gregorii; tropical East Africa), Buckollia (2; tropical East Africa), Camptocarpus (9; Madagascar, Mauritius), Cryptolepis (c 15; tropical and southern Africa, Madagascar, tropical Asia), Cryptostegia (2; Madagascar), Decalepis (4; India), Ectadium (3; the Namib Desert in Namibia), Epistemma (3; Ghana, Ivory Coast), Finlaysonia (3; tropical Asia and eastwards to tropical Australia), Gongylosperma (2; the Malay Peninsula), Gymnanthera (2; Malesia and eastwards to northern Australia), Hemidesmus (1; H. indicus; southern India, Southeast Asia, Malesia), Ischnolepis (1; I. graminifolia; Madagascar), Maclaudia (1; M. felixii; Guinea), Mangenotia (2; tropical West Africa), Meladerma (3; Thailand), Mondia (2; tropical and southern Africa), Myriopteron (1; M. extensum; northeastern India, southern China, Southeast Asia, West Malesia), Pentanura (2; Burma, Sumatra), Pentopetia (21; Madagascar), Periploca (14; the Mediterranean, North and tropical Africa, East Asia), Petopentia (1; P. natalensis; Eastern Cape, KwaZulu-Natal), Raphionacme (c 35; tropical and southern Africa, the Arabian Peninsula), Sacleuxia (2; tropical East Africa), Sarcorrhiza (1; S. epiphytica; tropical Africa), Schlechterella (2; tropical East Africa), Stelmacrypton (1; S. khasianum; eastern India, southern China), Stomatostemma (2; Zambia, Mozambique, Zimbabwe), Streptocaulon (9; tropical Asia), Tacazzea (4; tropical and southern Africa, southern Asia), Telectadium (3; Southeast Asia), Zacateza (1; Z. pedicellata; tropical Africa), Zygostelma (1; Z. benthamii; Thailand). – Tropical, subtropical and arid temperate regions of the Old World and eastwards to northern Australia. Roots often tuberous. Corolla sometimes with valvate aestivation. Nectariferous disc absent. Stamens inserted in corolla mouth. Staminal bases sometimes upright, connate and forming tube surrounding ovary. Nectaries present on margins of staminal bases and alternating with stamens. Anthers without lignified basal nectar guide appendages. Corpusculum basal. Pollen grains 4-16-porate, shed in calymmated (T-shaped tetragonal) tetrads, or in 20 pollinia (with inner walls reduced, free from translator apparatus), collected on translator (consisting of spoon-shaped structure + basal sticky viscidium). Exine smooth. Retinaculum formed through cellular fusion. Exotestal cells thickened or non-thickened. x = 11. – Phyllanthera may be sister to the remaining Periplocoideae.

[Apocyneae+[New World clade+[Baisseeae+[Secamonoideae+Asclepiadoideae]]]]

Apocyneae Rchb., Fl. Germ. Excurs. 1(3): 410, 429. Jul-Dec 1831

19/c 116. Papuechites (1; P. aambe; the Moluccas, New Guinea), Anodendron (17; tropical Asia to Japan and Vanuatu), Sindechites (1; S. henryi; China), Amphineurion (1; A. marginatum; northeastern India, southern China, Southeast Asia, Malesia), Streptoechites (1; S. chinensis; Hainan, Thailand, northern Vietnam), Vallaris (3; tropical Asia), Beaumontia (9; India and China and eastwards to Bali), Cleghornia (4; Sri Lanka, Southeast Asia, West Malesia), Apocynum (9; eastern Europe and southern Russia to China, temperate North America), Parameria (1; P. laevigata; India, southern China, Southeast Asia, Malesia), Aganonerion (1; A. polymorphum; Southeast Asia), Urceola (16; tropical Asia), Chonemorpha (10; tropical Asia), Trachelospermum (c 10; India to Japan, southeastern United States), Micrechites (12; tropical Asia to southern China and New Guinea), Amalocalyx (3; Southeast Asia), Pottsia (3; tropical Asia eastwards to Java), Aganosma (7; Southeast Asia, West and Central Malesia), Epigynum (7; Kashmir, Himalayas, southern China, tropical Asia and eastwards to tropical Australia). – Russia to East and tropical Asia, Vanuatu, North America. Often lianas. Pollen sometimes shed in tetrats. Stylar head sometimes with strap-shaped bands of adhesive. – Apocyneae are sister-group to the “New World clade”, according to Livshultz (2010). - Unplaced Apocyneae Baharuia (1; B. gracilis; Sumatra, Borneo), Eucorymbia (1; E. alba; Borneo), Ixodonerium (1; I. annamense; Southeast Asia), Parepigynum (1; P. funingense; Yunnan), Vallariopsis (1; V. lancifolia; West Malesia).

[New World clade+[Baisseeae+[Secamonoideae+Asclepiadoideae]]]

New World clade

c 28/415–430. The topology [Echiteae+[Mesechiteae+Odontadenieae]] is provisional and not very strongly supported.

Echiteae Bartl., Ord. Nat. Plant.: 204. Sep 1830 [‘Echitea’].

c 14/185–195. Laubertia (3; tropical America); Prestonia pro parte, Temnadenia (3; tropical South America), Peltastes (6–7; tropical America); ‘Echites’ (9; the West Indies; polyphyletic), ‘Prestonia’ (c 55; tropical America; paraphyletic), Artia (4–7; China, Southeast Asia, New Caledonia), Parsonsia (80–85; East Asia to Taiwan, tropical Asia and eastwards to northern Australia, New Caledonia, New Zealand, Fiji); unplaced: Asketanthera (4; tropical America), Ecua (1; E. moluccensis; the Moluccas), Hylaea (2; northeastern South America), Macropharynx (5; tropical South America), Thenardia (3–4; Central America), Thoreauea (3; Mexico). – Tropical America, East and tropical Asia and eastwards to northern Australia, New Caledonia, New Zealand, Fiji. – Laubertia may be sister to the remaining Echiteae.

[Mesechiteae+Odontadenieae]

Mesechiteae Miers, Apocyn. S. Amer.: 10. Mai-Jun 1878.

8/c 200. Elytropus (1; E. chilensis; central and southern Chile, Rio Negro in Argentina), Allomarkgrafia (11; Nicaragua to Peru), Forsteronia (c 40; tropical America), Mesechites (12; tropical America), Mandevilla (c 130; tropical America), Tintinnabularia (3; southern Mexico, Honduras), Pentalinon (2; Florida, Central America, the West Indies), Angadenia (2; Florida, the West Indies). – Florida, southern Mexico, Central America, the West Indies, tropical South America.

Odontadenieae Miers, Apocyn. S. Amer.: 8. Mai-Jun 1878.

6/30–35. Cycladenia (1; C. humilis; southwestern United States), Pinochia (4; Mexico, Central America, the Greater Antilles), Thyrsanthella (1; T. difformis; southeastern United States), Stipecoma (1; S. peltigera; Brazil), Odontadenia (c 20; tropical America), Secondatia (4–7; tropical America). – Southwestern United States, Mexico, tropical America. – Odontadenieae are not monophyletic in some of the analyses by Livshultz (2010).

[Baisseeae+[Secamonoideae+Asclepiadoideae]]

Baisseeae (Pichon ex De Kruif) M. E. Endress in Ann. Missouri Bot. Gard. 94(2): 266. Jul 2007 

4/29. Dewevrella (1; D. cochliostema; tropical Africa); Baissea (18; tropical and southern Africa), Motandra (3; tropical West and Central Africa), Oncinotis (7; tropical and southern Africa, Madagascar). – Tropical and southern Africa, Madagascar. Lianas. Hairs sometimes branched. Domatia sometimes present. Pollinia absent in Baissea. Stylar head sometimes with strap-shaped bands of adhesive. – Baisseeae are sister to [Secamonoideae+Asclepiadoideae]. Dewevrella is sister to the clade [Oncinotis+[Baissea+Motandra]].

[Secamonoideae+Asclepiadoideae]

Stamens inserted well below bases of corolla lobes. Corona usually formed from androecium. Nectariferous disc absent. Filaments absent. Gynostegium developed by postgenital fusion of androecial and gynoecial structures. Staminal feet erect, connate, forming tube around ovary. Anthers inserted on top of basal tube consisting of annular corona of staminal bases. Nectaries consisting of outgrowths of petaloid staminal bases (tissue from corolla and filaments) present behind nectar guides and alternating with stamens. Microsporogenesis successive. Pollen grains inaperturate, shed as T-shaped tetragonal tetrads. Pollen aggregated into pollinia in erect pollinaria. Pollinia of a pollinarium consisting of anther halves of adjacent stamens. Exine psilate. Translator – retinaculum – formed from hardened resinous mostly stigmatic secretion and apical adhesive corpusculum. Endosperm development usually nuclear (rarely cellular). Embryo with chlorophyll. Inulin and fructans sometimes present. Monoterpene indole alkaloids absent.

Secamonoideae Endl., Gen. Plant.: 589. Aug 1838 [‘Secamoneae’]

7–8/c 150. Calyptranthera (2; Madagascar), Genianthus (c 20; India, Bhutan and southern China to Malaysia and the Philippines), Goniostemma (2; India, China; in Toxocarpus?), Pervillea (4; Madagascar), Secamone (c 80; tropical and subtropical Africa, Madagascar, tropical and subtropical Asia and eastwards to northern Australia), Secamonopsis (2; Madagascar), Toxocarpus (c 40; tropical regions in the Old World eastwards to islands in the Pacific; incl. Goniostemma?), Trichosandra (1; T. borbonica; Mauritius). – Tropical and subtropical regions in the Old World eastwards to northern Australia and islands in the Pacific. Lianas. Corolla sinistrorsely or dextrorsely contorted. Pollinia 20, with outer walls reduced, connected to translator apparatus consisting of corpusculum (rarely also of one or two translator arms – caudicula). Pollen grains shed as tetrads. Exine with thick granular infratectal layer. x = 11.

Asclepiadoideae Burnett, Outlines Bot.: 1012, 1095, 1103. Feb 1835 [‘Stapelidae’, or ‘Asclepiadeae’]

145–150/2.430–2.560. Cosmopolitan except polar and cold-temperate regions, with their largest diversity in drier parts of Africa. Intraxylary phloem sometimes present. Leaves rarely alternate. Corolla often with valvate aestivation. Corona consisting of staminal parts. Anthers disporangiate (one sporangium from each theca absent). Nectaries present adjacent to or near base of nectar guides. Pollen tetrads linear during development. Pollinia ten (one pollinium from each theca fertile), with inner walls reduced, connected to translator apparatus made of corpusculum and (one or) two caudicula. Anther usually secreting wall material around pollinium. Compitum sometimes absent. Exine with thin granular infratectal layer. x = (9–)11(–14). - Unplaced Asclepiadoideae Vietnamia (1; V. inflexa; Vietnam).

[Fockeeae+[Marsdenieae+Ceropegieae+Asclepiadeae]]

Fockeeae H. Kunze, Meve et Liede in Taxon 43: 373. 31 Aug 1994

2/8. Fockea (6; tropical and southern Africa), Cibirhiza (2; Tanzania, Zambia, Oman). – Tropical and southern Africa, Oman. Pollen grains shed as tetrads. Pollinia erect, without sterile wall. Caudicula absent. – Fockeeae are sister to the remaining Asclepiadoideae.

[Marsdenieae+Ceropegieae+Asclepiadeae]

Pollinia with caudicula. Pollen grains shed as monads.

Marsdenieae Benth., Fl. Austral. 4: 325, 333. 16 Dec 1868

25/430–500. Anatropanthus (1; A. borneensis; Borneo), Anisopus (2; tropical West and Central Africa), Asterostemma (1; A. repandum; Java), Campestigma (1; C. purpureum; Southeast Asia), Clemensiella (1; C. mariae; the Philippines), Cosmostigma (3; Hainan, tropical Asia), Dischidia (c 80; northeastern India and southern China to Malesia, northeastern Queensland, New Caledonia), Dolichopetalum (1; D. kwangsiense; southern China), Gongronema (15; tropical regions in the Old World), Gunnessia (1; G. pepo; northernmost Queensland), Heynella (1; H. lactea; Java), Hoya (>200; tropical Asia, tropical and eastern Australia, islands in the Pacific to Polynesia), Lygisma (3; Southeast Asia), Marsdenia (>200; northern to southern Africa, Madagascar, tropical and subtropical Asia eastwards to Malesia, Australia, Melanesia, Central and South America, one species, M. erecta, in eastern Mediterranean), Oreosparte (1; O. celebica; Sulawesi), Pycnorhachis (1; P. maingayi; the Malay Peninsula), Rhyssolobium (1; R. dumosum; Namibia, Northern Cape), Sarcolobus (14; tropical Asia, tropical Australia, islands in western Pacific), Sphaerocodon (2; Zambia to Namibia), Spirella (2; Southeast Asia), Stigmatorhynchus (2–3; eastern and southern Africa), Telosma (10; tropical and southern Africa, Madagascar, tropical Asia eastwards to the Malay Peninsula), Treutlera (1; T. insignis; eastern Himalayas). – Tropical and subtropical regions on both hemispheres. Latex watery (clear). Pollinia erect or pendent with two caudicula. Sterile wall sometimes present on external side of pollinium. Exotestal cells in Hoya with external walls unthickened.

Ceropegieae Orb., Dict. Univ. Hist. Nat. 3: 339. 1 Jul 1843 [‘Ceropegiae’]

c 40/740–790. ‘Orbea’ (c 55; tropical and southern Africa, southwestern Arabian Peninsula; polyphyletic), ‘Tromotriche’ (11; southwestern Namibia, Northern, Western and Eastern Cape; polyphyletic), ‘Duvalia’ (23; southern Africa, Somalia, the Arabian Peninsula; polyphyletic), Piaranthus (8; Namibia, South Africa, Botswana), Australluma (2; Namibia, Zimbabwe and Mozambique to South Africa), Tridentea (8; southern Africa), ‘Stapelia’ (47; tropical and southern Africa; polyphyletic), Huernia (67; tropical and southern Africa, southern Arabian Peninsula), Tavaresia (3; tropical and southern Africa), Stapelianthus (9; Madagascar), Larryleachia (10; Namibia, South Africa), Richtersveldia (1; R. columnaris; Northern Cap), Notechidnopsis (2; Namaqualand in Northern and Western Cape), Hoodia (14; southwestern tropical and southern Africa), Lavrania (1; L. haagnerae; arid regions in Namibia and Northern Cape), Ophionella (1; O. arcuata; Eastern Cape), Baynesia (1; B. lophophora; northwestern Namibia), Pectinaria (3; Northern, Western and Eastern Cape), Stapeliopsis (6; Namibia, South Africa), Quaqua (c 30; Namibia, western South Africa), Duvaliandra (1; D. dioscoridis; Socotra), Socotrella (1; S. dolichocnema; Socotra), Whitesloanea (1; W. crassa; Somalia), ‘Caralluma’ pro parte, Rhytidocaulon (10; northeastern tropical Africa, the Arabian Peninsula), Echidnopsis (32; tropical East Africa, the Arabian Peninsula), Anomalluma (2; Somalia, southern Arabian Peninsula), ‘Caralluma’ pro parte (50–55; the Canary Islands, the Mediterranean, Africa, southwestern and southern Asia to Burma), Pseudolithos (6; northeastern tropical Africa; in Caralluma?), Edithcolea (1; E. grandis; eastern and northeastern Africa, the Arabian Peninsula, Socotra; in Caralluma?). – Unplaced Ceropegieae Anisotoma (2; Eastern Cape, KwaZulu-Natal), Brachystelma (>120; Africa, India and eastwards to New Guinea and tropical Australia), ‘Ceropegia’ (160–200; the Canary Islands, Africa, Madagascar, the Arabian Peninsula, India, Sri Lanka, southern China, Malesia eastwards to New Guinea and Queensland; non-monophyletic), Conomitra (1; C. linearis; Sudan), Dittoceras (3; eastern Himalayas, Thailand), Emplectanthus (2; KwaZulu-Natal), Heterostemma (12; tropical Asia, tropical Australia to eastern Queensland, islands in western Pacific), Leptadenia (4; tropical regions in the Old World), Neoschumannia (1; N. kamerunensis; tropical West and Central Africa), Orthanthera (4; southern tropical and southern Africa, India, Nepal), Pentasacme (4; tropical Asia), Riocreuxia (8–9; tropical and southern Africa, India), Sisyranthus (12; tropical and southern Africa). – The Canary Islands, the Mediterranean, tropical and subtropical regions in the Old World eastwards to eastern Queensland, islands in western Pacific, with their highest diversity in arid regions of eastern and southern Africa. Latex watery (clear). Pollinia erect with two caudicula, sterile wall present on apex or inner side of pollininium. Orbicules present in Riocreuxia.

Asclepiadeae Duby, Bot. Gall. 1: 323. 12-14 Apr 1828

c 85/1.360–1.370. Eustegia (5; E. minuta; Northern and Western Cape); Oncinema (1; O. lineare; Western and Eastern Cape), Microloma (11; Namibia, Northern, Western and Eastern Cape, Free State), Astephanus (2; southern Africa), Petalostelma (7; tropical South America), Minaria (21; tropical South America, with their highest diversity in the Espinhaço Range in eastern Brazil), Barjonia (6; Brazil), ‘Nephradenia’ (10; tropical America; paraphyletic), ‘Hemipogon’ (10; South America; polyphyletic), ‘Blepharodon’ (c 45; Central and South America, one species in North America; non-monophyletic), Peplonia (6; Brazil), Ditassa (c 55; South America), Tassadia (c 20; tropical America), Macroscepis (7–8; tropical America), Schubertia (6; South America), Gonolobus (c 150; tropical and subtropical America), ‘Matelea’ (c 180; tropical South America; non-monophyletic), Funastrum (16; United States, tropical America), Oxypetalum (c 135; tropical America, southern South America), Tweedia (6; tropical South America), Araujia (10–11; tropical South America), Philibertia (c 40; tropical South America), Pentacyphus (5; northern South America), Diplolepis (6; Chile, Argentina), Monsanima (2; tropical South America), Scyphostelma (10–15; tropical South America), Jobinia (3–4; tropical South America), Orthosia (c 20; tropical America), ‘Cynanchum’ (c 400; tropical and subtropical regions on both hemispheres; non-monophyletic), Glossonema (5; tropical and subtropical East Africa, the Arabian Peninsula, Pakistan), Pentarrhinum (2–3; tropical and southern Africa), Graphistemma (1; G. pictum; Hong Kong), Metaplexis (6; East Asia), Seshagiria (1; S. sahyadrica; western India), Holostemma (2; India, Sri Lanka, southern China, Southeast Asia), Raphistemma (2; tropical Asia), Schizostephanus (2; southern Africa to Angola and Zimbabwe), Pentatropis (5; tropical and subtropical regions in the Old World eastwards to Australia), Tylophora (c 50; tropical and subtropical regions of the Old World; paraphyletic), Vincetoxicum (c 70; Europe, temperate Asia, one species, V. carnosum, in Australia; incl. Tylophora?), Solenostemma (1; S. argel; Egypt, the Arabian Peninsula), Calciphila (2; Somalia), Oxystelma (2; tropical regions in the Old World), Pergularia (2; Africa, Madagascar and eastwards to India), Kanahia (2; tropical East Africa, the Arabian Peninsula), Calotropis (3; tropical and subtropical Africa and eastwards to India), ‘Asclepias’ (c 70; North America, Mexico, Central America; polyphyletic), Gomphocarpus (25–32; tropical and subtropical Africa and Asia), Xysmalobium (10–15; tropical and southern Africa), Aspidoglossum (c 35; tropical and southern Africa). – Unplaced Asclepiadeae Adelostemma (3; southern China, Burma), Aidomene (1; A. parvula; Angola), Amblystigma (7; Bolivia, Argentina), Anomotassa (1; A. macranthus; Ecuador), Aspidonepsis (5; Drakensberg in South Africa), Biondia (13; China), Blyttia (2; East Africa, southern Arabian Peninsula), Cordylogyne (1; C. globosa; southern Africa), Cyathostelma (2; Brazil), Diplostigma (1; D. canescens; East Africa), Emicocarpus (1; E. fissifolius; southeastern Africa), Fanninia (1; F. caloglossa; Eastern Cape to KwaZulu-Natal), Fischeria (16; tropical America), Glossostelma (12; tropical and southern Africa), Goydera (1; G. somaliensis; Somalia), Lagoa (1; L. calcarata; Brazil), Mahawoa (1; M. montana; Sulawesi), Margaretta (1; M. rosea; tropical Africa), Melinia (8; South America), Merrillanthus (1; M. hainanensis; Hainan), Miraglossum (7; eastern parts of southern Africa), Mitostigma (c 20; South America), Nautonia (1; N. nummularia; southern Brazil), Odontanthera (1; O. radians; Somalia, the Arabian Peninsula, southwestern Asia), Odontostelma (1; O. welwitschii; southern tropical Africa), Pachycarpus (37; tropical and southern Africa), Parapodium (3; southern Africa), Pentastelma (1; P. auritum; Hainan), Periglossum (4; southeastern and southern Africa), Pherotrichis (2; Mexico), Prosthecidiscus (1; P. guatemalensis; Central America), Rhyncharrhena (1; R. linearis; Australia), Rhyssostelma (1; R. nigricans; Argentina), Schizoglossum (c 15; tropical and southern Africa), Sichuania (1; S. alterniloba; China), Stathmostelma (13; eastern tropical Africa), Stenomeria (3; Colombia, Venezuela, Peru), Stenostelma (4–5; Namibia, South Africa, Botswana), Trachycalymma (10; tropical Africa), Trichosacme (1; T. lanata; Mexico), Vailia (2; South America), Vincetoxicopsis (1; V. harmandii; Southeast Asia), Widgrenia (1; W. corymbosa; Brazil), Woodia (3; Eastern Cape to KwaZulu-Natal and Mpumalanga). – Nearly cosmopolitan. Latex milky. Pollinia erect or pendant with two caudicula and with a sterile margin on top or on external side of the pollinium. – Eustegia minuta was recovered as sister to the remaining Asclepiadeae by Surveswaran & al. (2014).

Cladogram (simplified) of Apocynaceae based on DNA sequence data (Livshultz & al. 2007; Lens & al. 2008).

Stict consensus tree (simplified) of part of Apocynaceae based on DNA sequence data (Livshultz 2010).

GELSEMIACEAE (G. Don) Struwe et V. A. Albert

( Back to Rubiales )

Struwe et Albert in Cladistics 10: 206. Jun 1995

Pteleocarpaceae R. K. Brummitt in Kew Bull. 66: 3. Mar? 2011

Genera/species 3/12

Distribution Tropical Africa, Madagascar, southern China, Southeast Asia, West Malesia, southeastern United States to northeastern South America.

Fossils Unknown.

Habit Bisexual, evergreen tree (Pteleocarpa), shrubs or lianas (Gelsemium, Mostuea).

Vegetative anatomy Phellogen? Primary medullary strands narrow or wide. Secondary lateral growth anomalous (via a cylindrical cambium). Vessel elements usually with simple (sometimes scalariform or reticulate) perforation plates (vessel elements in Pteleocarpa solitary); lateral pits alternate, bordered pits. Vestured pits present in Pteleocarpa. Imperforate tracheary xylem elements tracheids (in Gelsemium) or fibre tracheids with simple and/or bordered pits, septate or non-septate. Wood rays biseriate (Pteleocarpa) to multiseriate, homocellular or heterocellular. Axial parenchyma absent or rare (paratracheal scanty, unilateral or banded); in Pteleocarpa usually apotracheal diffuse-in-aggregates or unilateral banded. Intraxylary phloem present. Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace. Parenchyma in Pteleocarpa often with gum-like substances. Styloids, druses and elongate calciumoxalate crystals present.

Trichomes Hairs unicellular or absent.

Leaves Usually opposite (sometimes verticillate, in Pteleocarpa alternate, spiral), simple, entire, with convolute ptyxis (Pteleocarpa). Stipules two, interpetiolar to intrapetiolar or somewhat decurrent, or absent; leaf sheath absent. Axillary colleters present (especially in connection to stipules). Petiole bases sometimes connate via an ochrea or a line. Petiole vascular bundle transection arcuate? Venation pinnate, brochidodromous. Stomata paracytic or anomocytic. Cuticular waxes? Domatia as pockets or absent. Leaf margin faintly sinuate or entire.

Inflorescence Terminal or axillary, cymose (often panicle; flowers sometimes solitary).

Flowers Usually actinomorphic (sometimes slightly zygomorphic). Hypogyny. Sepals usually five (sometimes four), with imbricate quincuncial aestivation, sometimes unequal in length, persistent, free (Gelsemium) or connate in lower part (Mostuea, Pteleocarpa). Petals usually five (sometimes four), with imbricate quincuncial aestivation, sometimes unequal in length, connate into an infundibuliform or tubular perigone. Nectary? Disc absent. Heterostyly often present.

Androecium Stamens usually five (sometimes four), haplostemonous, antesepalous, alternipetalous. Filaments free, adnate to corolla tube (epipetalous). Anthers dorsifixed, versatile, tetrasporangiate, extrorse (Gelsemium, Pteleocarpa) or latrorse (Mostuea), longicidal (dehiscing by longitudinal splits). Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolporate, shed as monads, ?-cellular at dispersal. Exine tectate, with columellate infratectum, striate, finely reticulate or punctate.

Gynoecium Pistil composed of two connate carpels. Ovary superior, bilocular. Stylodia two, connate in lower part, stylar branches often bifid. Stigmas two or four, capitate, type? Pistillodium absent.

Ovules Placentation axile. Ovules two (Mostuea, Pteleocarpa) or two to eight (Gelsemium) per carpel (in Pteleocarpa one large pendulous epitropous and one small orthotropous and later aborted ovule), anatropous or amphitropous?, unitegmic, tenuinucellar. Integument ? cell layers thick. Megagametophyte monosporous, Polygonum type? Endosperm development nuclear? Endosperm haustoria? Embryogenesis?

Fruit In Gelsemium and Mostuea a loculicidal and/or septicidal capsule with persistent calyx; in Pteleocarpa a large flat suborbicular one-seeded samara with one wide membranous wing on each side, divided by suture leading to retuse apex.

Seeds Aril absent. Seed flattened. Testa thin, winged or hairy. Exotestal cells? Endotesta? Perisperm not developed. Endosperm copious, horny, starchy (Gelsemium). Embryo small, straight or curved, chlorophyll? Cotyledons two. Germination?

Cytology n = 8, 10

DNA

Phytochemistry Flavonols (kaempferol, quercetin), O-methylated flavones, Route I iridoids (also secoiridoids), Group IX secoiridoids (indole alkaloids of corynanthe type), and C-17 indole alkaloids present. Caffeic acid?

Use Ornamental plants, medicinal plants, timber (Pteleocarpa).

Systematics Mostuea (8; tropical Africa, Madagascar, northeastern South America), Gelsemium (3; southern China, Southeast Asia, West Malesia, southern United States, eastern Mexico, Guatemala), Pteleocarpa (1; P. lamponga; West Malesia)

Gelsemiaceae are probably sister-group to Apocynaceae.

Nuclear ribosomal ETS analysis suggests Pteleocarpa being sister to [Gelsemium+Mostuea], whereas Mostuea is sister to [Gelsemium+Pteleocarpa] in analyses of plastid regions (Struwe & al. 2014).

The leaves are spirally arranged in Pteleocarpa, not opposite as in the majority of Gentianales. Gottwald (1982) suggested a possible affinity with Apocynaceae and Rubiaceae judging from wood anatomical similarities. However, Pteleocarpa is most closely related to Gelsemium and Mostuea, according to molecular analyses (Miller 2003; Struwe & al. 2014).

GENTIANACEAE Juss.

( Back to Rubiales )

de Jussieu, Gen. Plant.: 141. 4 Aug 1789 [’Gentianae’], nom. cons.

Coutoubeaceae Martinov, Tekhno-Bot. Slovar: 168. 3 Aug 1820 [‘Coutoubeae’]; Obolariaceae Martinov, Tekhno-Bot. Slovar: 427. 3 Aug 1820 [’Obolariae’]; Chironiaceae Bercht. et J. Presl, Přir. Rostlin 1(104-114): 1, 36. 1823 [‘Chironieae’]; Potaliaceae Mart., Nov. Gen. Sp. Plant. 2: 89, 91, 133. Jan-Jun 1827 [‘Potalieae’]; Chironiineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 1071, 1078. 1846 [‘Chironieae’]; Gentianineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 1071. 1846 [‘Gentianeae’]; Chironiales Griseb., Grundr. Syst. Bot.: 144. 1-2 Jun 1854 [‘Chironiflorae’]; Saccifoliaceae Maguire et Pires in Mem. New York Bot. Gard. 29: 242. 14 Jun 1978; Voyriaceae Doweld, New Syllabus Pl. Fam.: 857. Apr 2007

Genera/species c 90/1.600–1.650

Distribution Cosmopolitan except polar and arid areas, with their largest diversity in temperate and subtropical regions and on tropical mountains.

Fossils Unknown.

Habit Usually bisexual (rarely polygamomonoecious), perennial, biennial or annual herbs (sometimes shrubs, rarely lianas or trees). Some genera are partially mycoheterotrophic. A few clades (Cotylanthera, Leiphaimos, Voyria, and Voyriella) consisting of achlorophyllous holoparasitic mycotrophs.

Vegetative anatomy Mycorrhiza often Paris type. Phellogen ab initio superficial. Primary medullary strands narrow. Medulla often with vascular bundles. Endodermis often prominent. Parenchyma often septate. Secondary lateral growth normal or anomalous, from cylindrical cambium. Vessel elements with usually simple (in Saccifolium scalariform) perforation plates; lateral pits alternate, simple or bordered pits. Vestured pits present. Imperforate tracheary xylem elements libriform fibres with simple and/or bordered pits, usually non-septate. Wood rays usually uniseriate, homocellular or heterocellular, or absent. Axial parenchyma apotracheal diffuse?, or paratracheal scanty vasicentric, scalariform or banded (absent in Saccifolium). Intraxylary phloem present. Sieve tube plastids S type. Nodes ≥1:≥1, at least unilacunar with one or more leaf traces (trilacunar nodes common, sometimes multilacunar, with several traces; sometimes with split lateral traces). Medulla sometimes with sclereids. Acicular crystals, styloids, crystal sand and other types of calciumoxalate crystals present in some species.

Trichomes Hairs unicellular or multicellular, simple, uniseriate or multiseriate, or absent; glandular hairs sometimes present.

Leaves Usually opposite (rarely verticillate; in Saccifolium alternate, spiral), simple, entire, sometimes coriaceous (in mycotrophs scale-like, reduced), with varying ptyxis (in Saccifolium saccate-vaginate near apex). Stipules absent (interpetiolar or intrapetiolar sheath-like or auriculate structures sometimes present at nodes); leaf sheath absent. Axillary colleters present in some species (in association with stipules). Leaves often connate at base. Petiole vascular bundles? Venation usually palmate (sometimes pinnate; in Saccifolium appearing parallelodromous). Stomata usually anomocytic (sometimes anisocytic). Cuticular wax crystalloids? Epidermis and/or mesophyll with or without mucilage cells. Mesophyll sometimes with sclerenchymatous idioblasts. Leaf margin usually entire (sometimes crenate). Extrafloral nectaries sometimes present on leaves (e.g. in Anthocleista and Irlbachia).

Inflorescence Terminal or axillary, usually dichasial, sometimes panicle or fasciculate cymose (rarely racemose; flowers sometimes solitary).

Flowers Usually actinomorphic (rarely zygomorphic). Epicalyx present or absent. Hypogyny. Sepals usually four or five (rarely up to 16), usually with imbricate (sometimes valvate or open) aestivation, often persistent, usually connate (rarely free). Petals usually four or five (rarely up to 16), usually with contorted (rarely imbricate) aestivation, marcescent, more or less connate; scales or nectariferous pits often present inside corolla tube (corolla tube in Halenia with five spurs). Nectariferous glands often present. Disc intrastaminal, annular or composed of separate parts, or absent. Diheterostyly present in some species.

Androecium Stamens usually four or five (rarely up to 16), antesepalous, alternipetalous. Filaments usually free (sometimes connate in lower part into a tube), adnate to corolla tube (epipetalous). Anthers usually free (sometimes connate at base; rarely connate into a tube), usually dorsifixed and versatile (sometimes basifixed and non-versatile), tetrasporangiate, usually introrse (rarely extrorse or latrorse), usually longicidal (dehiscing by longitudinal slits; in, e.g., Exacum poricidal, with apical pores; anthers rarely septate); connective sometimes slightly prolonged (sometimes glanduliferous); placentoids sometimes present. Tapetum usually secretory (sometimes amoeboid-periplasmodial), with uninucleate cells. Staminodia one to four, alternating with fertile stamens, or absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually (2–)3(–4)-colporate or (1–)2–3-porate, usually shed as monads (sometimes as tetrads), bicellular or tricellular at dispersal. Exine tectate or semitectate, with columellate infratectum, perforate, reticulate, microreticulate, striate, echinulate, verrucate, regulate, or psilate. Pollen tubes sometimes several.

Gynoecium Pistil composed of two connate carpels (carpels median or transverse). Ovary superior, usually unilocular (rarely bilocular; rarely divided by secondary septa), sometimes with a gynophore. Style usually single, simple (sometimes persistent; absent in Lomatogonium). Stigma usually widely bilobate (sometimes capitate or subpeltate; in Lomatogonium decurrent along sides of ovary), papillate, Wet type. Pistillodium absent.

Ovules Placentation usually intrusively parietal (rarely basal or axile; in Cotylanthera free central). Ovules several to numerous per carpel, usually anatropous (sometimes hemianatropous or orthotropous), horizontal, usually unitegmic (in some mycotrophic species [Exacum, Voyria] ategmic), tenuinucellar (reduced, with meiocyte semi-inferior). Integument two to 20 cell layers thick. Hypostase present. Megagametophyte monosporous, Polygonum type (in some mycoheterotrophic species at least seemingly inverted 180o). Antipodal cells sometimes proliferating (diploid to polyploid, sometimes multiplying, persistent; in Halenia multinucleate). Endosperm development usually nuclear (in Voyriella cellular). Endosperm haustoria? Embryogenesis solanad.

Fruit Usually a septicidal capsule (sometimes a berry), often with persistent calyx.

Seeds Seeds sometimes winged. Aril absent. Testa multiplicative, exotestal. Exotestal cells sometimes elongate, usually with thickened inner walls. Endotesta disintegrating. Perisperm not developed. Endosperm usually copious (sometimes sparse), oily (starch absent). Embryo small, straight, well differentiated or rudimentary (in Cotylanthera, Leiphaimos, Voyria and Voyriella 5- to 24-celled, undifferentiated), with or without chlorophyll. Cotyledons two (rarely much reduced). Germination phanerocotylar.

Cytology n = 5<, 9–15, 17 etc.

DNA Deletion of 100 bp present in plastid gene trnL.

Phytochemistry Flavonols, O-methylated flavones, flavone-O-glucosides (in two species of Exacum), flavone-C-glycosides (in Potalieae and Gentianeae), Route I iridoids, Group VI secoiridoids (secologanin), Group VII secoiridoids (sweroside, swertiamarin, gentiopicroside, gentianin), Group X secoiridoids (loganin, iridoid pyridine alkaloids), caffeic acid, alkaloids, saponins, cyanogenic compounds, and 6-substituted xanthones (norathyriol etc., in Chironieae and Gentianeae) present. Indole alkaloids? Ellagic acid, tannins and proanthocyanidins not found. Carbohydrates stored as oligosaccharides (not starch). Aluminium accumulated in a few species.

Use Ornamental plants, medicinal plants.

Systematics Gentianaceae are sister-group to the [Loganiaceae+[Gelsemiaceae+Apocynaceae]] clade. A plausible topology is the following (Merckx & al. 2013): [Saccifolieae+[Exaceae+[Voyrieae+[Chironieae+[Potalieae+[Helieae+Gentianeae]]]]]]

Saccifolieae (Maguire et Pires) Struwe, Thiv, V. A. Albert & Kadereit in L. Struwe et V. A. Albert, Gentianac.: Syst. Nat. Hast.: 48. 2002

5/16–20. Curtia (6–10; Guianas to Uruguay), Hockinia (1; H. montana; eastern Brazil), Saccifolium (1; S. bandeirae; Guayana Highlands), Tapeinostemon (7; northeastern South America), Voyriella (1; northeastern South America). – Northeastern South America to Uruguay. Sometimes shrubs or achlorophyllous mycoheterotrophic herbs. Leaves sometimes alternate (spiral). Flowers usually pentamerous (sometimes tetra- or hexamerous). Heterostyly sometimes present. Placentation parietal. Endosperm development in Voyriella cellular. Cotyledons sometimes absent. n = 10–14.

[Exaceae+[Voyrieae+[Chironieae+[Potalieae+[Helieae+Gentianeae]]]]]

Exaceae Colla, Herb. Pedem. 4: 174. 15-31 Aug 1835

8/145–150. Lagenias (1; L. pusillus; Western Cape), Sebaea (c 40; tropical and subtropical Africa, Madagascar, Socotra, the Arabian Peninsula and eastwards to India and Sri Lanka, southern Australia, Tasmania, New Zealand), Exochaenium (22; tropical and southeastern Africa), Exacum (c 70; tropical regions in the Old World, with their highest diversity in Madagascar), Klackenbergia (2; Madagascar), Ornichia (3; Madagascar), Gentianothamnus (1; G. madagascariensis; Madagascar), Tachiadenus (11; Madagascar). – Tropical regions in the Old World, southern Australia, Tasmania, New Zealand, with their highest diversity in Madagascar. Some species of Exacum and Sebaea achlorophyllous mycoheterotrophic herbs. Flowers usually actinomorphic (in Exacum and Orphium zygomorphic), usually pentamerous (sometimes tetramerous). Sepals free or connate. Sebaea with one pair of collateral secondary stigmas at stylar base. Ovules in some species of Exacum orthotropous, ategmic. Anticlinal exotestal cell walls sinuate. n = 9, 11, 15 or more. Flavone-O-glycosides present in few species of Exacum.

[Voyrieae+[Chironieae+[Potalieae+[Helieae+Gentianeae]]]]

Voyrieae Gilg in H. G. A. Engler et K. A. E. Prantl, Nat. Pflanzenfam. IV, 2: 62, 102. Jun 1895

1/19. Voyria (19; tropical America, one species, V. primuloides, in tropical Africa). – Achlorophyllous mycoheterotrophic herbs. Roots and shoots exogenous or endogenous. Rhizoids usually absent. Vascular bundles bicollateral, separate. Colleters present or absent. Pollen grains 1–6-porate. Exine scabrate to smooth. Sometimes with stamen-like appendages (staminodia or nectaries?) on each side of ovary. Stigma infundibular. Ovules orthotropous ategmic or anatropous unitegmic. Endothelium present. Nucellar cap present. Endosperm development cellular or nuclear. Fruit a septicidal capsule. Seed coat exotestal. Endosperm sparse to almost absent. Embryo undifferentiated. n = 16–20.

[Chironieae+[Potalieae+[Helieae+Gentianeae]]]

Placentation parietal. Xanthones and L-(+)-bornesitol present.

Chironieae Dumort., Fl. Belg.: 51. 1827

27/170–180. Bisgoeppertia (2; the West Indies), Blackstonia (4; Europe, the Mediterranean), Centaurium (c 20; Europe, the Mediterranean, temperate Asia), Chironia (c 30; tropical and southern Africa, Madagascar), Cicendia (2; C. filiformis: Europe, the Mediterranean; C. quadrangularis: Oregon, California, western South America), Zeltnera (c 25; southwestern United States, northwestern Mexico), Gyrandra (3; Mexico, Central America), Schenkia (5; Europe, the Mediterranean, Australia, Tasmania, the Hawaiian Islands), Eustoma (3; southern United States and Mexico to northern South America), Exaculum (1; E. pusillum; Europe), Geniostemon (4; Mexico), Ixanthus (1; I. viscosus; the Canary Islands), Orphium (1; O. frutescens; Western Cape), Sabatia (17–20; North America, Mexico, Central America, the West Indies), Zygostigma (2; Brazil, Argentina); Canscora (9; tropical regions in the Old World), Cracosna (3; Southeast Asia), Duplipetala (2; Southeast Asia, West Malesia), Hoppea (2; India, Sri Lanka, Burma), Microrphium (1; M. pubescens; West Malesia to the Philippines), Phyllocyclus (5; southern China, Burma), Schinziella (1; S. tetragona; tropical Africa); Coutoubea (5; the West Indies, tropical South America), Deianira (7; tropical America), Schultesia (15–20; tropical America), Symphyllophyton (2; Brazil), Xestaea (1; X. lisianthoides; Panamá, Venezuela). – Subcosmopolitan, with their highest diversity in tropical and subtropical regions. Usually chlorophyllous herbs (sometimes shrubs). Flowers (2–)4–5(–12)-merous. Sepals connate. Pollen grains sometimes shed as tetrads. n = 10, 13–15, 17 or more. Special 6-substituted xanthones.

[Potalieae+[Helieae+Gentianeae]]

Nectaries present.

Potalieae Endl., Gen. Plant.: 576. Aug 1838

13/c 155. Congolanthus (1; C. longidens; tropical Africa), Djaloniella (1; D. ypsilostyla; the Diaguissa Plateau in Guinea), Enicostema (3; tropical Africa and Madagascar and eastwards to the Lesser Sunda Islands, Central America, the West Indies), Faroa (17–19; tropical Africa), Karina (1; K. tayloriana; Congo), Lisianthius (c 30; tropical America), Neurotheca (3; tropical Africa, one species, N. loeselioides, also in tropical South America), Oreonesion (1; O. testui; Gabon), Pycnosphaera (1; P. buchananii; tropical Africa), Urogentias (1; U. ulugurensis; Uluguru and Nguru Mountains in Tanzania); Anthocleista (c 50; tropical Africa, Madagascar, the Mascarene Islands), Fagraea (60–70; southern India, Sri Lanka and eastwards to China, Southeast Asia, Malesia and islands in the western Pacific, with their highest diversity on Borneo), Potalia (9; tropical America). – Pantropical, with their highest diversity in Africa, Madagascar and Malesia. Trees, shrubs, lianas, herbs. Nodes ≥5:≥5, multilacunar with at least five leaf traces. Epidermal and cortical sclereids often present. Interpetiolar or intrapetiolar sheath-like structures or auricles often present. Flowers 3–16(–24)-merous. Sepals connate at base. Carpellary fusion sometimes congenital. Fruit sometimes a berry. n = ? Flavone-C-glucosides present. – Potaliinae: nodes multilacunar, sclereids present, flowers up to 16-merous (Anthocleista, Potalia), corolla caducous, pollen grains porate, carpellary fusion congenital, fruit a berry. Anthocleista has more than two metres long seedling leaves.

[Helieae+Gentianeae]

Helieae Gilg in Engler et Prantl, Nat. Pflanzenfam. IV, 2: 62, 95. Jun 1895

17/c 195. Aripuana (1; A. cullmaniorum; Amazonian Brazil), Celiantha (3; Guayana Highlands), Chorisepalum (5; Guayana Highlands), Irlbachia (17; tropical South America), Lehmanniella (4; Panamá, Colombia, Peru), Macrocarpaea (c 105; tropical America), Neblinantha (2; Guayana Highlands), Prepusa (5; mountains in Brazil), Rogersonanthus (3; northeastern South America, Trinidad), Roraimaea (1; R. coccinea; southern Venezuela, northern Brazil), Senaea (2; mountains in Brazil), Sipapoantha (2; southern Venezuela, northern Brazil), Symbolanthus (c 30; tropical America), Tachia (12; tropical South America), Tetrapollinia (1; T. caerulescens; Venezuela, Guianas), Wurdackanthus (2; the West Indies, northeastern South America), Yanomamua (1; Y. araca; Amazonian Brazil), Zonanthus (1; Z. cubensis; Cuba). – Tropical America, with their largest diversity in northeastern South America. Usually herbs (sometimes shrubs). Interpetiolar or intrapetiolar stipules or sheath-like structure sometimes present. Flowers usually pentamerous (sometimes tetra- or hexamerous). Pollen grains sometimes shed as tetrads or polyads. Style often long, spirally twisted and flattened as dry. n = ?

Gentianeae Dumort., Anal. Fam. Plant.: 25. 1829

c 20/890–920. Gentiana (c 350; temperate and arctic-alpine regions on both hemispheres, tropical mountains), Kuepferia (12; northern India, Sikkim, southern and eastern Tibet, northwestern Yunnan, southwestern Sichuan, Nepal, Bhutan, northern Burma), Crawfurdia (16; northern India, Sikkim, Bhutan, China, northern Burma), Metagentiana (14; China, northern Burma, northern Thailand), Sinogentiana (2; China), Tripterospermum (24–25; East Asia); Bartonia (3–4; eastern North America), Comastoma (c 25; temperate and arctic-alpine regions on the Northern Hemisphere, the Himalayas), Frasera (15; North America), ‘Gentianella’ (c 200; temperate regions on the Northern Hemisphere, eastern New South Wales, Victoria, Tasmania, the Andes; polyphyletic), Gentianopsis (16–25; temperate regions on the Northern Hemisphere), Halenia (c 80; mountain regions in Europe, Asia and North America, the Andes), Jaeschkea (3–4; Himalayas), Latouchea (1; L. fokienensis; eastern China), ’Lomatogonium’ (18–21; temperate and arctic-alpine regions in Europe and Asia; polyphyletic), Megacodon (1; M. stylophorus; Himalayas), Obolaria (1; O. virginica; eastern North America), Pterygocalyx (1; P. volubilis; East Asia), Lomatogoniopsis (3; China), ’Swertia’ (135–150; Europe, temperate Asia, Madagascar, mountain regions in Africa and Malesia; polyphyletic), Veratrilla (2; eastern Himalayas, western China). – Northern temperate, to Sulawesi (some species of Tripterospermum), and Africa and Madagascar (some species of Swertia). Corolla sometimes with nectaries. Style often short or absent. n = 5 or higher, very variable. Special xanthones, flavone-C-glucosides present. Fructans or inulin sometimes present.

Cladogram (simplified) of Gentianaceae based on DNA sequence data (Merckx & al. 2013).

LOGANIACEAE R. Br. ex Mart.

( Back to Rubiales )

von Martius, Nov. Gen. Sp. Plant. 2: 133. Jan-Jun 1827 [’Loganieae’], nom. cons.

Strychnaceae DC. ex Perleb, Vers. Artzneikr. Pfl.: 244. Mai 1818 [’Strychneae’]; Spigeliaceae Bercht. et J. Presl, Přir. Rostlin 1: 40. 1823 [’Spigelieae’]; Strychnales Link, Handbuch 1: 439. 4-11 Jul 1829 [‘Strychnaceae’]; Loganiales Lindl., Nix. Plant.: 19. 17 Sep 1833; Gardneriaceae Wall. ex Perleb, Clav. Class.: 23. Jan-Mar 1838 [’Gardnereae’]; Spigeliineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 1071, 1081. 1846 [‘Spiegelieae’]; Strychnineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 1051, 1060. 1846 [‘Strychneae’]; Antoniaceae Hutch., Fam. Fl. Pl., ed. 2: 375. 4 Jun 1959; Geniostomaceae L. Struwe et V. A. Albert in Cladistics 10: 206. Jun 1995

Genera/species 13/c 415

Distribution Tropical and subtropical regions in both hemispheres.

Fossils Unknown.

Habit Usually bisexual (rarely monoecious, gynomonoecious, dioecious, or gynodioecious), evergreen or deciduous trees, shrubs or lianas (with branch tendrils), perennial or annual herbs.

Vegetative anatomy Phellogen ab initio superficially or deeply seated. Primary medullary strands usually narrow (sometimes wide or alternately narrow and wide). Secondary lateral growth normal or anomalous (from cylindrical cambium). Vessel elements usually with simple (rarely scalariform or reticulate) perforation plates; lateral pits alternate, bordered pits. Vestured pits sometimes present (Geniostoma). Imperforate tracheary xylem elements 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 paratracheal scanty vasicentric, confluent or banded, or absent. Intraxylary phloem (diffuse) present in, e.g., Strychnos. Tyloses sometimes frequent. Sieve tube plastids S type. Nodes 1:≥1, unilacunar with one or more leaf traces, 3:3, trilacunar with three traces, or multilacunar with several traces (sometimes with split lateral strands). Prismatic calciumoxalate crystals often abundant; styloids present in some species.

Trichomes Hairs usually unicellular simple (sometimes multicellular, multi-armed, stellate, candelabra-shaped or lepidote; sometimes glandular) or absent.

Leaves Usually opposite (rarely verticillate or pseudoverticillate), simple, entire, sometimes coriaceous (rarely scale-like), often with flat ptyxis. Stipules interpetiolar, intrapetiolar (sometimes sheathing) or absent; leaf sheath absent. Axillary colleters present in many species (especially in association with stipules). Petiole bases usually connate, often via a line or short ochrea. Petiole vascular bundles? Venation usually pinnate (sometimes palmate; leaves sometimes one-veined). Stomata anomocytic, paracytic, anisocytic or cyclocytic. Cuticular wax crystalloids? Domatia as pockets or hair tufts or absent. Epidermis with or without mucilage cells. Mesophyll with or without sclerenchymatous idioblasts (branched stone cells). Leaf margin serrate or entire. Extrafloral nectaries present or absent.

Inflorescence Terminal or axillary, dichasial, panicle, thyrsoid, spike-like, corymb or umbel-like cymose (in Mitreola and Spigelia sometimes spike-like cincinni; flowers sometimes solitary axillary). Some species of Mitrasacme with pseudanthia at fruit ripening surrounded by accrescent involucre consisting of bracts or possibly floral prophylls (bracteoles).

Flowers Usually actinomorphic (in Usteria zygomorphic). Hypogyny (rarely partial epigyny). Sepals (four or) five, with imbricate, valvate or open aestivation, uniform or non-uniform (rarely petaloid), often persistent, usually connate at base (median sepal in Logania abaxial), sometimes with adaxial colleters. Petals (four or) five, usually with imbricate or valvate (rarely contorted) aestivation, connate into an infundibuliform or tubular corolla, often with adaxial hairs at lobe bases. Nectariferous disc poorly developed, intrastaminal, or absent. Nectaries sometimes present on ovary walls. Diheterostyly present in some genera (e.g. Geniostoma).

Androecium Stamens usually five (sometimes four; in Usteria one, abaxial), haplostemonous, antesepalous, alternipetalous. Filaments free, adnate to corolla tube (epipetalous). Anthers usually dorsifixed (sometimes basifixed), versatile?, tetrasporangiate, usually introrse (sometimes latrorse or extrorse), longicidal (dehiscing by longitudinal slits); connective sometimes slightly prolonged at apex. Tapetum secretory. Staminodia one to three, extrastaminal, or absent; female flowers often with staminodia. Secondary pollen display present in some species.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually (2–)3(–4)-colporate (rarely 2–4-colpate, tripororate, polypantoporate, polyzonoporate or syncolpate), shed as monads, tricellular at dispersal. Exine tectate or semitectate, with columellate infratectum, perforate to reticulate, psilate.

Gynoecium Pistil composed of two (or three) usually connate carpels (sometimes partially free; carpels sometimes congenitally syncarpous; sometimes postgenitally connate exclusively at apex). Ovary usually superior (rarely semi-inferior), usually bilocular (rarely unilocular or trilocular). Style single, simple, thick, or stylodia two (rarely three), free (stylodia rarely connate at apex only), often persistent, or absent. Stigma capitate, punctate, clavate, truncate, or bilobate (or trilobate), papillate, Wet type (Mitreola). Male flowers often with pistillodium.

Ovules Placentation usually axile (rarely apical). Ovules usually several to numerous (rarely one) per carpel, usually anatropous or hemianatropous (rarely amphitropous, campylotropous or orthotropous), unitegmic, tenuinucellar. Integument four to six cell layers thick. Endothelium present in Mitrasacme. Megagametophyte monosporous, Polygonum type. Endosperm development usually nuclear (rarely cellular). Endosperm haustoria lateral or absent. Embryogenesis usually solanad (in Strychnos and allied taxa onagrad).

Fruit A loculicidal and/or septicidal or denticidal capsule, a follicle (rarely a pyxidium) or a schizocarp with two follicle-like mericarps; in Strychnos, Neuburgia etc. a drupe or a berry.

Seeds Aril absent. Testa thin or thick, sometimes winged or (in Geniostoma and Labordia) embedded in juicy tissue or swollen placentae. Exotestal cells hairy or papillate, thick-walled and lignified (except outer wall). Endotesta? Perisperm not developed. Endosperm usually copious (rarely sparse), carnose or horny (rarely ruminate), with starch or hemicellulose. Embryo large or small, straight or curved, well differentiated, chlorophyll? Cotyledons two. Germination phanerocotylar.

Cytology n = 10–12, 16

DNA Mitochondrial coxI intron present in Strychnos.

Phytochemistry Flavonols (kaempferol, quercetin), O-methylated flavones, Route I iridoids, Group VI secoiridoids (secologanin), Group VII secoiridoids (sweroside, gentiopicroside), Group IX secoiridoids (indole alkaloids of corynanthe-type), Group X secoiridoids (desoxyloganin, loganin, ketologanin), tryptophane-derived alkaloids, indole alkaloids (in Strychneae), and saponins present. Ellagic acid, proanthocyanidins and cyanogenic compounds not found. Caffeic acid? Aluminium accumulated in some species of Strychnos and allied groups.

Use Ornamental plants, medicinal plants, poisons (Strychnos), fruits (Strychnos spp.), timber.

Systematics Usteria (1; U. guineensis; tropical West and Central Africa), Bonyunia (c 10; Colombia to Brazil, Peru and Bolivia), Antonia (1; A. ovata; tropical South America); Gardneria (5; India, central Japan and southwards to Southeast Asia and Java), Neuburgia (10–12; the Philippines and Sulawesi to New Guinea, New Caledonia, Vanuatu and Fiji), Spigelia (c 50; tropical and subtropical America), Strychnos (c 190; tropical and subtropical regions on both hemispheres), Mitrasacme (c 55; East and tropical Asia, Australia, Tasmania, New Caledonia, New Zealand), Orianthera (13; Australia, with their highest diversity in southwestern Western Australia), Geniostoma (c 40; the Mascarene Islands, Malesia to southern Japan, Bonin Islands and Taiwan, eastern Queensland, Vanuatu, Fiji, New Zealand, Samoa, Tonga, Tahiti, Henderson Island, the Hawaiian Islands), Mitreola (6; tropical regions on both hemispheres), Logania (c 22; Australia, New Caledonia, New Zealand, with their highest diversity in southwestern Western Australia, South Australia and New South Wales); Norrisia (2; West Malesia).

Loganiaceae are sister to [Gelsemiaceae+Apocynaceae].

Consensus tree of Loganiaceae from successive weighting based on DNA sequence data (Backlund & al. 2000).

RUBIACEAE Juss.

( Back to Rubiales )

de Jussieu, Gen. Plant.: 196. 4 Aug 1789, nom. cons.

Cinchonaceae Batsch, Tab. Affin. Regni Veg.: 234. 2 Mai 1802 [’Cinchoneae’]; Coffeaceae Batsch, Tab. Affin. Regni Veg.: 233. 2 Mai 1802; Guettardaceae Batsch, Tab. Affin. Regni Veg.: 235. 2 Mai 1802 [’Guettardeae’]; Aparinaceae Hoffmanns. et Link, Fl. Portug. 2: 3, 38. 1813-1829 [’Aparines’]; Operculariaceae Juss. ex Perleb, Vers. Artzneikr. Pfl.: 207. Mai 1818 [’Operculariae’]; Spermacocaceae Bercht. et J. Presl, Přir. Rostlin: 256. Jan-Apr 1820 [’Spermacoceae’]; Catesbaeaceae Martinov, Tekhno-Bot. Slovar: 111. 3 Aug 1820 [’Catesbiaceae’]; Coutareaceae Martinov, Tekhno-Bot. Slovar: 168. 3 Aug 1820 [’Coutareae’]; Hydrophylacaceae Martinov, Tekhno-Bot. Slovar: 318. 3 Aug 1820 [’Hydrophylaceae’]; Nonateliaceae Martinov, Tekhno-Bot. Slovar: 420. 3 Aug 1820 [’Nonateliae’]; Pagameaceae Martinov, Tekhno-Bot. Slovar: 447. 3 Aug 1820 [’Pagameae’]; Randiaceae Martinov, Tekhno-Bot. Slovar: 534. 3 Aug 1820 [’Randiae’]; Sabiceaceae Martinov, Tekhno-Bot. Slovar: 555. 3 Aug 1820 [’Sabiceae’]; Cephalanthaceae Raf. in Ann. Gén. Sci. Phys. Bruxelles 6: 86. Oct-Dec 1820 [’Cephalantia’]; Rubiineae Raf. in Ann. Gén. Sci. Phys. Bruxelles 6: 83. Oct-Dec 1820 [‘Rubiacea‘]; Hedyotidaceae Dumort., Comment. Bot.: 57. Nov-Dec 1822 [’Hediotideae’]; Gardeniaceae Dumort., Anal. Fam. Plant.: 29, 32. 1829; Theligonaceae Dumort., Anal. Fam. Plant.: 15, 17. 1829 [’Theligoneae’], nom. cons.; Lygodisodeaceae Bartl., Ord. Nat. Plant.: 123, 207. Sep 1830 [’Lygodysodeaceae’]; Psychotriaceae F. Rudolphi, Syst. Orb. Veg.: 49. 5-12 Jul 1830 [’Psychotrieae’]; Rubiopsida Bartl., Ord. Nat. Plant.: 123, 206. Sep 1830 [’Rubiacinae’]; Cinchonales Lindl., Nix. Plant.: 18. 17 Sep 1833; Asperulaceae Cham. ex Spenn., Handb. Angew. Bot. 2: 492. 1835 [’Asperuleae’]; Hameliaceae Mart., Consp. Regn. Veg.: 31. Sep-Oct 1835 [‘Hamelieae’]; Lygodisodeales Bartl. in C. F. P. von Martius, Consp. Regn. Veg.: 31. Sep-Oct 1835 [‘Lygodyseaceae’]; Galiaceae Lindl., Intr. Nat. Syst. Bot., ed. 2: 249. 13 Jun 1836 [’Stellateae, or Galiaceae’]; Galiales Bromhead in Edinburgh New Philos. J. 24: 414. Apr 1838; Lippayaceae Meisn., Plant. Vasc. Gen.: Tab. Diagn. 156, Comm. 112. 16-22 Sept 1838 [’Lippayeae’]; Houstoniaceae Raf., Good Book: 21. Jan 1840 [’Houstonides’]; Cynocrambaceae Meisn., Plant. Vasc. Gen., Tab. Diagn. 345, Comm. 256. 13-15 Feb 1842 [’Cynocrambeae’], nom. illeg.; Coffeopsida Brongn., Enum. Plant. Mus. Paris: xvii, 48. 12 Aug 1843 [’Coffeineae’]; Cinchonineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 790, 791. 1846 [‘Cinchoneae’]; Coffeineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 791, 826. 1846 [‘Coffeaceae’]; Gardeniineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 791, 804. 1846; Guettardineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 791, 819. 1846; Hameliineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 791, 817. 1846; Hedyotidineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 791, 812. 1846; Spermacocineae J. Presl in Nowočeská Bibl. [Wšobecný Rostl.] 7: 791, 835. 1846 [‘Spermacoceae’]; Naucleaceae (DC.) Wernh. in New Phytol. 11: 225. 30 Jul 1912; Theligonales Nakai in J. Jap. Bot. 18: 92, 94. 10 Mar 1942 [’Thelygonales’]; Dialypetalanthaceae Rizzini et Occhioni in Lilloa 17: 253. 30 Dec 1948, nom. cons.; Henriqueziaceae Bremek. in Acta Bot. Neerl. 6 [Meded. Bot. Mus. Herb. Rijksuniv. Utrecht 141]: 371. Aug 1957

Genera/species 535–550/11.470–11.550

Distribution Cosmopolitan except polar areas, with their highest diversity in tropical and subtropical regions.

Fossils Fossil pollen grains of Rubiaceae have been found from the Late Eocene onwards. Fruits of Emmenopterys are known from mid-Eocene layers in Oregon and Washington. Canthium and Guettarda are reported from the Late Eocene of Australia and Faramea from the Late Eocene of Panamá. Fossil leaves were described as Paleorubiaceophyllum from mid-Eocene layers in Tennessee and Kentucky and further genera are known from the Oligocene and, above all, the Miocene (Graham 2009). Cephalanthus is recorded from Cenozoic strata in Europe.

Habit Usually bisexual (rarely monoecious, polygamomonoecious or dioecious), evergreen or deciduous trees, shrubs, lianas or suffrutices, or perennial, biennial or annual herbs. Hydnophytum, Myrmecodia, Myrmeconauclea, Myrmephytum, etc. are epiphytic or (Myrmeconauclea) rheophytic myrmecophytes (ant plants), in the hollow swollen stems and branches – hypocotylar bases – of which ant colonies live. Young stems and branches often quadrangular in cross-section.

Vegetative anatomy Phellogen ab initio usually superficial (sometimes deeply seated). Primary medullary strands at least usually narrow. Secondary lateral growth usually normal (sometimes anomalous from concentric cambia). Endodermis sometimes significant. Vessel elements usually with simple (sometimes scalariform or reticulate) perforation plates; lateral pits alternate, bordered pits. Vestured pits present. Imperforate tracheary elements tracheids or fibre tracheids with simple and/or bordered pits, usually non-septate. Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty vasicentric, confluent, aliform-confluent, reticulate or banded, or absent. Tyloses sometimes abundant. Intraxylary phloem absent. Sieve tube plastids Ss type. Nodes 1(–≥3):1(–≥3), usually unilacunar (rarely trilacunar to multilacunar) with one leaf trace (rarely several traces), lateral bundles splitting off and forming vascular cylinder around stem and innervating stipules. Parenchyma often with secretory cells and cavities with gum-like substances. Colleters of different types often present. Prismatic crystals, crystal sand, druses, styloids, acicular and other (calciumoxalate?) crystal types often present; idioblasts (raphid cells) with calciumoxalate raphides present in, e.g., Rubioideae.

Trichomes Hairs unicellular or multicellular, simple or branched, cylindrical, papillose, moniliform, T-shaped, spiral or band-shaped (sometimes stellate or lepidote), in Rubioideae often septate; glandular hairs often present.

Leaves Usually opposite (sometimes verticillate), simple, usually entire (in Genipa and Posoqueria lobed), usually with flat ptyxis. Stipules usually interpetiolar (innervated from vascular cylinder), often serrate (sometimes intrapetiolar, interpetiolar-intrapetiolar or sheathing), often connate and adnate to petioles, caducous or persistent; leaf sheath absent. Axillary colleters abundant (especially in connection with stipules). Petiole vascular bundle transection arcuate or annular. Venation pinnate (leaves sometimes one-veined). Stomata usually paracytic (sometimes parallelocytic). Cuticular waxes? Domatia as pockets or hair tufts or absent (bacteriodomatia with Burkholderia strains present in some species of Pavetta, Psychotria and Sericanthe). Mesophyll with or without sclerenchymatous idioblasts. Calciumoxalate raphides present in Rubioideae; druses occurring in many species lacking raphides. Leaf margin serrate or entire. Extrafloral nectaries present on lamina in some species (e.g. in Tocoyena).

Inflorescence Terminal or axillary, panicle, thyrse, thyrsoid, pseudoverticillate, capitate, etc. (flowers sometimes solitary axillary). Some species with capitate pseudanthia bearing involucre consisting of (sometimes petaloid) bracts.

Flowers Usually actinomorphic (in Gleasonia, Henriquezia and Platycarpum zygomorphic). Epicalyx sometimes present. Usually epigyny (sometimes half epigyny; in Gaertnera and Pagamea secondary hypogyny; in Theligonum?). Sepals (three or) four or five, usually with open (sometimes imbricate, contorted or valvate) aestivation, often persistent (sometimes accrescent in fruit), usually connate (sometimes very small; rarely large and petaloid; in Dialypetalanthus 2+2, free; absent in Theligonum?). Petals (three or) four or five (to ten), with imbricate, valvate or contorted (rarely cochleate) aestivation, usually connate into hypocrateriform, tubular or infundibuliform corolla (in Dialypetalanthus 2+2, antesepalous, free; absent in Theligonum?). Nectariferous disc fleshy, annular (rarely absent). Diheterostyly often present.

Androecium Stamens (three or) four or five, antesepalous, alternipetalous (in Dialypetalanthus [eight to] 16 to 17 [to 25] in two whorls; in Theligonum [two to] seven to twelve [to 30], sometimes basally fused in fascicles of two, four or six stamens). Filaments usually free (in Dialypetalanthus connate at base), usually adnate to corolla tube (epipetalous; not in Dialypetalanthus). Anthers usually free (in, e.g., Argostemma connate), basifixed or dorsifixed, sometimes versatile, tetrasporangiate, usually introrse (sometimes extrorse), usually longicidal (dehiscing by longitudinal slits; in many genera poricidal, with apical pits); connective sometimes prolonged. Tapetum secretory. Staminodia absent. Secondary pollen display present in numerous Ixoroideae.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually colporate (sometimes colpate, porate or inaperturate; in Theligonum 4–8-zonoporate), usually shed as monads (sometimes as tetrads), bicellular or tricellular at dispersal. Exine tectate or semitectate, with columellate infratectum, perforate, reticulate or microreticulate, spinulate, echinate, verrucate or with other types of supratectal processes, or psilate, or intectate.

Gynoecium Pistil composed of two (to five; rarely up to 16) connate carpels (in Theligonum seemingly? monocarpellate, with a pseudomonomerous? gynoecium). Ovary usually inferior (sometimes semi-inferior, rarely secondarily superior), usually bilocular (rarely unilocular, trilocular, quadrilocular, quinquelocular, or up to novemlocular). Style single and narrow, or stylodia two (to five) and free or partly connate (style in Theligonum gynobasic); often with apical or subapical thickening, on which pollen grains are deposited (secondary pollen display). Stigma capitate, or usually bilobate (rarely trilobate to quinquelobate), papillate or non-papillate, Dry or Wet type. Pistillodium absent.

Ovules Placentation usually axile (sometimes apical, rarely parietal [when unilocular] or basal). Ovule one (usually in Rubioideae) to numerous per carpel, usually anatropous or hemitropous or campylotropous? (rarely circinotropous or hypertropous; in Theligonum campylotropous or amphitropous), pendulous, horizontal or ascending, apotropous or epitropous, unitegmic, usually tenuinucellar (reduced, with meiocyte semi-inferior; rarely crassinucellar; megasporangium sometimes absent). Integument one to 14 cell layers thick. Funicular obturator often present. Megasporocytes often several. Parietal cells sometimes present. Megasporangial epidermal cells sometimes anticlinally elongate. Megagametophyte usually monosporous, Polygonum type (rarely tetrasporous, Drusa type, or disporous, Allium type; megagametophytes sometimes several, haustorial). Antipodal cell(s) sometimes persistent, occasionally proliferating and haustorial. Endosperm development usually nuclear (rarely cellular). Endosperm haustoria? Embryogenesis usually ? (sometimes solanad). Suspensor haustorium present in some species.

Fruit A loculicidal and/or septicidal capsule (rarely a pyxidium), a berry or a drupe (sometimes a schizocarp with two [to nine] nut-like mericarps, sometimes a pseudofruit consisting of fused ovaries; pericarp in Theligonum with elaiosome).

Seeds Seeds sometimes pachychalazal. Aril absent? Only exotesta persistent (testa sometimes absent); exotesta usually one-layered; tangential walls of exotestal cells often with thickenings or exotestal cells parenchyma-like (sometimes palisade-like). Mesotestal cell walls sometimes thickened. Endotesta usually parenchymatous and crushed. Perisperm not developed. Endosperm usually copious, horny or soft (sometimes ruminate), oily (often with hemicellulose, in Theligonum with starch; sometimes sparse). Embryo large or small, straight or curved, usually well differentiated, without chlorophyll. Cotyledons two, usually flat. Germination phanerocotylar or cryptocotylar.

Cytology n = 9–11(–17)

DNA atpB promoter usually lost in Rubioideae. Plastid gene infA lost/defunct (Galium, Pentas). Mitochondrial intron coxII.i3 lost (Coffea). Mitochondrial coxI intron present in many genera.

Phytochemistry Flavonols (kaempferol, quercetin), flavones, O-methylated flavones, Group I carbocyclic iridoids (geniposide, scandoside, daphylloside, monotropein, gardenoside), Group II carbocyclic iridoids (shantziside), Group IX secoiridoids (ipecacalkaloids, indole alkaloids of corynanthe type), Group X secoiridoids (loganin, antirrhide, gardoside), proanthocyanidins, quinoline alkaloids (e.g. quinine), iridoid coumarins (asperuloside), tannins, ursolic acid, caffeic acid, saponins, anthraquinones (usually shikimic acid derived, although in Asperula polyacetate derived, e.g. alizarin), and arbutin present. Cinchonoideae with corynantheane and complex indole alkaloids. Phenylalanine-derived cyanogenic compounds? Ellagic acid not found. Aluminium accumulated in, e.g., numerous species of Rubioideae.

Use Ornamental plants, medicinal plants, flavouring (quinine from Cinchona), perfumes (Gardenia etc.), stimulants (Coffea), beverages (Morinda), tanning, dyeing sources (Morinda, Rubia etc.), timber.

Systematics Luculia may be sister to all other Rubiaceae, and [Coptosapelta+Acranthera] successive sister to the remainder. However, in some analyses Luculia is recovered as sister to the clade [Coptosapelteae+Rubioideae] (see, e.g., Manns 2012). Cinchonoideae are probably sister-group to Ixoroideae.

Luculieae Rydin et B. Bremer in C. Rydin et al. in Plant Syst. Evol. 278: 120. Mar 2009

1/5. Luculia (5; Himalayas, Yunnan, Indochina, Malesia). – Shrubs or small trees. Inflorescence umbel or corymb. Pollen grains tricolporate, acolumellate(?). – Luculia is often member of a basal polytomy also comprising Coptosapelteae, a third branch consisting of [Cinchonoideae+Ixoroideae] and a fourth branch consisting of Rubioideae. On the other hand, Luculia may be sister to [Coptosapelteae+Rubioideae] (with fairly weak support).

Coptosapelteae Bremek. ex S. P. Darwin in Taxon 25: 598. 26 Nov 1976

2/c 48. Coptosapelta (13; southeastern China to Malesia), Acranthera (c 35; tropical Asia, with their highest diversity on Borneo). – India, China, Southeast Asia, Western and Central Malesia. Raphides present. Hairs T-shaped. Corolla with dextrorsely contorted aestivation. Pollen grains 3–5-pororate. Exine with acolumellate infratectum, reticulate to psilate. Diheterostyly or secondary pollen presentation occurring. Aluminium accumulated. – It is possible that Coptosapelteae are sister-group to Rubioideae (see, e.g., Manns & al. 2012).

[Cinchonoideae+Ixoroideae]

Cinchonoideae Raf. in Ann. Gén. Sci. Phys. Bruxelles 6: 81. Oct-Dec 1820 [‘Cinchonaria’]

c 103/1.500–1.530. Cosmopolitan, but mainly Neotropical. Woody. Hairs usually cylindrical. Raphides present in Hamelieae and Hillieae. Calycophylly present in some species. Corolla contorted (usually sinisterally) or cochleate. Secondary pollen presentation frequently occurring. Stigma usually bilobate. Ovules often numerous per carpel. Fruit usually a capsule. Route II iridoids (carboxylated iridoids) and indole alkaloids present. Corynanthean and complex indole alkaloids present in e.g. Cinchoneae. Aluminium accumulated in some species. – A possible topology is the following (Manns & al. 2012): [[Cinchoneae+Isertieae]+[[Hymenodictyoneae+Naucleeae]+[Guettardeae+Rondeletieae]+[Chiococceae+[[Chione+Colleteria]+[Hamelieae+Hillieae]]]]]

[Cinchoneae+Isertieae]

Cinchoneae DC. in Ann. Mus. Natl. Hist. Nat. 9: 217. 1807 [‘Cinchonaceae’]

9/110–120. Joosia (11; western South America), Stilpnophyllum (4; the Andes in Ecuador and Peru), Ciliosemina (2; northwestern South America), Ladenbergia (c 35; tropical America), Cinchonopsis (1; C. amazonica; central and western Amazonas), Cinchona (20–25; Costa Rica, Colombia and Venezuela to central Bolivia), Remijia (35–40; tropical South America), Maguireocharis (1; M. neblinae; Guayana Highlands)?, Pimentelia (1; P. glomerata; Peru)? – Tropical America. – Cinchoneae are sister to Isertieae.

Isertieae A. Rich. ex DC., Prodr. 4: 342, 435. late Sep 1830

2/15. Isertia (14; tropical America), Kerianthera (1; K. preclara; Amazonian Brazil). – Tropical America.

[[Hymenodictyoneae+Naucleeae]+[Guettardeae+Rondeletieae]+[Chiococceae+[[Chione+Colleteria]+[Hamelieae+Hillieae]]]]

[Hymenodictyoneae+Naucleeae]

Hymenodictyoneae Razafim. et B. Bremer in Syst. Geogr. Plants 71: 535. 2002

2/c 32. Paracorynanthe (2; Madagascar), ‘Hymenodictyon’ (c 30; tropical regions in the Old World eastwards to Sulawesi; paraphyletic). – Tropical regions in the Old World eastwards to Sulawesi. Corolla with valvate aestivation. – Hymenodictyoneae seem to be sister to Naucleeae.

Naucleeae Burnett, Outlines Bot.: 906. Feb 1835

17/180–200. Cephalanthus (6; tropical regions on both hemispheres, southeastern United States), Mitragyna (13; tropical regions in the Old World), Sinoadina (1; S. racemosa; China, Japan, Burma, Thailand), Haldina (1; H. cordifolia; India, Sri Lanka, southern China, Southeast Asia), Adina (11; China, the Korean Peninsula, Japan, tropical Asia to New Guinea), Khasiaclunea (1; K. oligocephala; northeastern India; in Adina?), Neonauclea (65–70; southern China, tropical Asia and eastwards to western Pacific islands), Diyaminauclea (1; D. zeylanica; Sri Lanka; in Neonauclea?), Breonadia (1; B. salicina; tropical Africa, Madagascar), ‘Breonia’ (6–20; Madagascar; paraphyletic), Gyrostipula (2; Madagascar, the Comoro Islands), Janotia (1; J. macrostipula; Madagascar), Corynanthe (10–15; tropical Africa), Pseudocinchona (2; tropical Africa), Uncaria (c 40; tropical regions on both hemispheres), Neolamarckia (2; tropical Asia and eastwards to tropical Australia), Nauclea (12; tropical regions in the Old World). – Tropical regions on both hemispheres, East Asia, one species in southeastern United States.

[Guettardeae+Rondeletieae]

Guettardeae DC. in Ann. Mus. Natl. Hist. Nat. 9: 217. 1807 [‘Guettardaceae’]

c 20/680–685. Rogiera (c 15; southern Mexico, Central America, the West Indies, northern South America), Allenanthus (3; Central America), Neoblakea (2; N. venezuelensis; Venezuela), ‘Antirhea’ (c 35; Madagascar, Malesia and eastwards to Samoa; polyphyletic), Bobea (4; the Hawaiian Islands), Hodgkinsonia (2; eastern Queensland, northeastern New South Wales; in Anthirhea?), Tinadendron (2; Vanuatu, New Caledonia; in Anthirhea?), ‘Guettarda’ (c 150; Vanuatu, New Caledonia, tropical America, one species, G. speciosa, along tropical coasts; polyphyletic), Timonius (c 170; Mauritius, the Seychelles, Sri Lanka, the Andaman Islands, Malesia and eastwards to islands in the Pacific, with their largest diversity on New Guinea), ‘Chomelia’ (20–25; tropical America; polyphyletic), ‘Arachnothryx‘(107; Mexico, Central America, Trinidad, Colombia to Peru; paraphyletic), Javorkaea (1; J. hondurensis; Honduras; in Arachnothryx?), Cuatrecasasiodendron (2; Colombia; in Arachnothryx?), Gonzalagunia (c 35; tropical America), Malanea (c 35; tropical America), ‘Machaonia’ (c 30; tropical America; polyphyletic), Dichilanthe (2; Sri Lanka, Borneo)?, Neolaugeria (35; the West Indies), Stenostomum (c 30; the West Indies, northern South America; incl. Resinanthus?), Resinanthus (16; the West Indies, one species, R. aromaticus, in Mexico, with their highest diversity on Cuba and Hispaniola; in Stenostomum?), Pittoniotis (2; tropical America). – Madagascar, Mauritius, the Seychelles, Sri Lanka, the Andamans, Malesia and eastwards to eastern Australia, Melanesia, Samoa and the Hawaiian Islands, tropical America. – Guettardeae are sister to Rondeletieae.

Rondeletieae Burnett, Outlines Bot.: 906. Feb 1835

c 12/90–95. ‘Rondeletia’ (45–50; Panamá, the West Indies, tropical South America; polyphyletic), Rovaeanthus (2; southern Mexico, Central America), Suberanthus (7; Cuba, Hispaniola), Acunaeanthus (1; A. tinifolius; Cuba), Blepharidium (2; Central America), Acrosynanthus (7; the West Indies), Roigella (1; R. correifolia; western Cuba), Arcythophyllum (c 15; tropical America), Phyllomelia (1; P. coronata; western Cuba; in Mazaea?), Mazaea (2; Cuba), Habroneuron (1; H. radicans; Mexico), Lindenia (2–3; L. austrocaledonica: New Caledonia; L. vitiensis: Fiji; ?L. rivalis: Central America). – Melanesia, tropical America.

[Chiococceae+[[Chione+Colleteria]+[Hamelieae+Hillieae]]]]

Chiococceae Benth. et Hook. f., Gen. Plant. 2: 9, 21. 7-9 Apr 1873

28/190–195. Strumpfia (1; S. maritima; the West Indies); Exostema (c 25; tropical America, with their highest diversity in the West Indies), Siemensia (1; S. pendula; western Cuba), Bikkia (c 20; East Malesia to islands in western Pacific), Erithalis (10; Florida, the West Indies), Badusa (3; Palawan, New Guinea, western Pacific islands), Scolosanthus (21; the West Indies), Chiococca (6; southern Florida, tropical America), Schmidtottia (16; eastern Cuba), Ceuthocarpus (1; C. involucratus; eastern Cuba), Salzmannia (1; S. nitida; eastern Brazil), Eosanthe (1; E. cubensis; eastern Cuba), Phialanthus (18; the West Indies), Ceratopyxis (1; C. verbenacea; western Cuba), Coutarea (7; Mexico, Central America to Argentina), Isidorea (c 20; the West Indies), Portlandia (6; Jamaica), Cubanola (2; Cuba, Hispaniola), Cigarrilla (1; C. mexicana; Mexico), Nernstia (1; N. mexicana; Mexico), Osa (1; O. pulchra; Costa Rica), Thogsennia (1; T. lindeniana; eastern Cuba, Hispaniola), Phyllacanthus (1; P. grisebachianus; western Cuba), Catesbaea (c 20; Florida Keys, the West Indies), Asemnantha (1; A. pubescens; Mexico), Coutaportla (2; Mexico), Morierina (2; New Caledonia), Shaferocharis (3; eastern Cuba). – East Malesia, Melanesia, tropical America, with their highest diversity in the West Indies.

[[Colleteria+Chione]+[Hillieae+Hamelieae]]

[Colleteria+Chione]

2/3. Chione (1; C. venosa; Central America, the West Indies), Colleteria (2; the West Indies). – Central America, the West Indies.

[Hillieae+Hamelieae]

Raphides present.

Hillieae Bremek. ex S. P. Darwin in Taxon 25: 603. 26 Nov 1976

3/29. Cosmibuena (4; tropical America), Balmea (1; B. stormae; Mexico), Hillia (24; tropical America); – Tropical America. – Hillieae sensu stricto (comprising Cosmibuena, Balmea and Hillia) are sister to Hamelieae.

Hamelieae A. Rich. ex DC., Prodr. 4: 342, 438. late Sep 1830

8/c 175. Cosmocalyx (1; C. spectabilis; Mexico), Deppea (c 25; central and southern Mexico, Central America, southeastern Brazil), Syringantha (1; S. coulteri; Mexico), Hamelia (c 40; tropical America), Pinarophyllon (2; Central America), Plocaniophyllon (1; P. flavum; Chiapas in Mexico), Omiltemia (4; Mexico), Hoffmannia (c 100; Mexico, Central America and southwards to Argentina). – Tropical America.

Ixoroideae Raf. in Ann. Gén. Sci. Phys. Bruxelles 6: 84. Oct-Dec 1820 [‘Ixorinia’]

220–225/3.700–3.740. Pantropical. Latex-like substance sometimes present. Corolla with contorted or cochleate aestivation. Stigma usually bilobated. Placentation usually axile (sometimes parietal). Fruit often a drupe or a berry. Embryo short. – A plausible topology is the following: [Posoquerieae+Sipaneeae]+Condamineeae+[[Mussaendeae+Sabiceeae]+[Steenisia+[Retiniphyllum+[“Vanguerieae alliance”+”Coffeeae alliance”]]]]

[Posoquerieae+Sipaneeae]

This clade may be sister-group to the remaining Ixoroideae.

Posoquerieae Delprete in P. G. Delprete et al., Fl. Ilustr. Catarin. Rubiaceas 1: 23. 27 Dec 2004

5/c 40. Gleasonia (5; tropical South America), Henriquezia (7; Amazonian Brazil), Platycarpum (12; northern South America), Molopanthera (1; M. paniculata; eastern Brazil), Posoqueria (15–16; tropical America). – Tropical America. Flowers in Gleasonia, Henriquezia and Platycarpum zygomorphic. – Posoquerieae are sister to Sipaneeae.

Sipaneeae Bremek. in Rec. Trav. Bot. Néerl. 31: 252. 1934

10?/c 50. Maguireothamnus (2; Venezuela), Sipanea (19; tropical South America), Neobertiera (1; N. gracilis; Guyana), Sipaneopsis (6; northwestern South America), Dendrosipanea (3; northern South America), Limnosipanea (7; Panamá to tropical South America), Chalepophyllum (5; Venezuela, Guyana), Steyermarkia (1; S. guatemalensis; Central America), Neblinathamnus (3; Venezuela)?, Pteridocalyx (2; Guyana)? – Central America, tropical South America.

Condamineeae Benth. et Hook. f., Gen. Plant. 2: 12. 7-9 Apr 1873

31/280–285. Dioicodendron (1; D. dioicum; northwestern tropical South America), Emmenopterys (1; E. henryi; southwestern China, Burma, Thailand), Pinckneya (1; P. bracteata; southeastern United States), Ferdinandusa (20–25; tropical America), Dolicholobium (c 30; the Philippines and eastwards to Fiji), Mussaendopsis (2; West Malesia), Mastixiodendron (7; East Malesia and eastwards to Fiji), Capirona (1; C. decorticans; northeastern South America), Parachimarrhis (1; P. breviloba; Amazonia), Simira (c 45; tropical America), Calycophyllum (c 10; tropical America), Alseis (15–20; southern Mexico, Central America to Brazil), Dialypetalanthus (1; D. fuscescens; eastern Brazil), Bothriospora (1; B. corymbosa; tropical America), Wittmackanthus (1; W. stanleyanus; tropical America), Dolichodelphys (1; D. chlorocrater; Colombia, Ecuador, Peru), Chimarrhis (13; tropical America), Warszewiczia (8; tropical America), Bathysa (c 15; Amazonia), Picardaea (2; Cuba, Hispaniola), Pogonopus (2–3; tropical America), Condaminea (3; the Andes), Macbrideina (1; M. peruviana; Ecuador, Peru), Elaeagia (c 15; tropical America), Macrocnemum (6–8; Central America, Colombia), Rustia (c 15; tropical America), Hippotis (12; tropical America), Pentagonia (c 35; tropical America), Sommera (12; tropical America), Tammsia (1; T. anomala; Colombia, Venezuela), Hintonia (4; Central America). – Southeast Asia, Malesia, Melanesia, southeastern United States, tropical America. – Condamineeae may be sister-group to the remaining Ixoroideae. Dialypetalanthus of eastern Brazil is sister to the clade [Bothriospora+Wittmackanthus]. The phloem is stratified, and the perianth members are decussate and free from each other. Moreover, Dialypetalanthus has (8–)16–17(–25) basally connate stamens with poricidal anthers.

[[Mussaendeae+Sabiceeae]+[Steenisia+[Retiniphyllum+[“Vanguerieae alliance”+”Coffeeae alliance”]]]]

[Mussaendeae+Sabiceeae]

Mussaendeae Benth. et Hook. f., Gen. Plant. 2: 8, 15. 7-9 Apr 1873

c 6/170–175. Heinsia (4–5; tropical Africa), Bremeria (c 20; Madagascar, the Mascarene Islands), Pseudomussaenda (6; tropical Africa), ’Mussaenda’ (c 140; tropical regions in the Old World; non-monophyletic), Landiopsis (1; L. capuronii; Madagascar), Neomussaenda (2; Borneo). – Tropical regions in the Old World (absent from Australia and eastwards). – Mussaenda is characterized by reduplicate-valvate aestivation and glabrous style, whereas Bremeria is distinguished from the remaining Mussaendeae by having reduplicate-valvate as well as induplicate-valvate aestivation, and densely pubescent style.

Sabiceeae A. Stahl, Estud. Fl. Puerto-Rico 5: 32. 1887 [‘Sabicieae’]

4/c 140. Sabicea (c 130; tropical Africa, Madagascar, tropical America), Tamridaea (1; T. capsulifera; Socotra), Virectaria (8; tropical Africa), Hekistocarpa (1; H. minutiflora; Nigeria, Cameroun). – Tropical Africa, Madagascar, Socotra, tropical America. – Sabiceeae seem to be sister-group to Mussaendeae and [Mussaendeae+Sabiceeae] in turn sister to [Steenisia+[Retiniphyllum+[Crossopteryx+[Scyphiphora+[Vanguerieae+[Greeneeae+[Alesanthieae+Ixoreae]]]]]]].

[Steenisia+[Retiniphyllum+[“Vanguerieae alliance”+”Coffeeae alliance”]]]

Steenisieae Kainul. et B. Bremer in K. Kainulainen, S. G. Razafimandimbison et B. Bremer, Bot. J. Linn. Soc. 173: 397. [DOI: 10.1111/boj.12038]. 25 Apr 2013

1/5. Steenisia (5; Borneo, Natuna Islands). – Corolla with contorted aestivation. – Steenisia is sister to the remaining Ixoroideae.

[Retiniphyllum+[Crossopteryx+[Scyphiphora+[Vanguerieae+[Greeneeae+[Alesanthieae+Ixoreae]]]]]]

Retiniphylleae Benth. et Hook. f., Gen. Plant. 2: 9, 20. 7-9 Apr 1873

1/c 20. Retiniphyllum (c 20; Colombia, Venezuela, Brazil, the eastern Andes). – Resiniferous. Corolla with contorted aestivation. Stamens with basal and apical sterile appendages. Ovary quinquelocular. Ovules two per locule, collateral, pendulous. Drupe one-seeded. – Retiniphylleae are sister-group to the [“Vanguerieae alliance”+”Coffeeae alliance”] clade.

Vanguerieae alliance”

The “Vanguerieae alliance” comprises Crossopteryx, Jackiopsis, Scyphiphora, Trailliaedoxa, Glionnetia, Vanguerieae, Greeneeae, Aleisanthieae, and Ixoreae.

[Crossopteryx+[Jackiopsis+[Scyphiphora+[Trailliaedoxa+[Glionnetia+[Vanguerieae+[Greeneeae+[Aleisanthieae+Ixoreae]]]]]]]]

Crossopterygeae F. White ex Bridson, Fl. Zambes. 5(3): 389. 2003

1/1. Crossopteryx (1; C. febrifuga; tropical and southern Africa). – Crossopteryx may be sister to the remaining “Vanguerieae alliance”.

[Jackiopsis+[Scyphiphora+[Trailliaedoxa+[Glionnetia+[Vanguerieae+[Greeneeae+[Aleisanthieae+Ixoreae]]]]]]]

Jackieae Korth. in Ned. Kruidk. Arch. 2/2): 196. 1851 [‘Jackiae’]

1/1. Jackiopsis (1; J. ornata; the Malay Peninsula, Borneo). – Jackiopsis may be sister to the remaining “Vanguerieae alliance”.

[Scyphiphora+[Trailliaedoxa+[Glionnetia+[Vanguerieae+[Greeneeae+[Aleisanthieae+Ixoreae]]]]]]

Scyphiphoreae Kainul. et B. Bremer in K. Kainulainen, S. G. Razafimandimbison et B. Bremer, Bot. J. Linn. Soc. 173: 397. [DOI: 10.1111/boj.12038]. 25 Apr 2013

1/1. Scyphiphora (1; S. hydrophyllacea; coastal areas in Southeast Asia, West Malesia, tropical Australia and New Caledonia). – Mangrove shrub. – Scyphiphora may be sister to the remaining “Vanguerieae alliance”.

[Trailliaedoxa+[Glionnetia+[Vanguerieae+[Greeneeae+[Aleisanthieae+Ixoreae]]]]]

Trailliaedoxeae Kainul. et B. Bremer in K. Kainulainen, S. G. Razafimandimbison et B. Bremer, Bot. J. Linn. Soc. 173: 397. [DOI: 10.1111/boj.12038]. 25 Apr 2013

1/1. Trailliaedoxa (1; T. gracilis; southwestern China). – Trailliaedoxa may be sister to the remaining “Vanguerieae alliance”.

[Glionnetia+[Vanguerieae+[Greeneeae+[Aleisanthieae+Ixoreae]]]]

Glionnetia

1/1. Glionnetia (1; G. sericea; the Seychelles). – Glionnetia may be sister to the clade [Vanguerieae+[Greeneeae+[Aleisanthieae+Ixoreae]]], although the support for this position is weak (see Kainulainen & al. 2013, where it was part of a trichotomy).

[Vanguerieae+[Greeneeae+[Aleisanthieae+Ixoreae]]]

Vanguerieae (A. Rich.) Dumort., Anal. Fam. Plant.: 32. 1829

c 23/590–600. Keetia (c 40; tropical and southern Africa), Psydrax (c 100; tropical regions in the Old World eastwards to tropical Australia and islands in the Pacific), Afrocanthium (c 20; eastern and southern Africa), ‘Pyrostria’ (c 35; tropical Africa, Madagascar, Mauritius, Rodrigues, the Seychelles; paraphyletic), Bullockia (8–9; eastern and southern Africa, Madagascar; in Pyrostria?), Peponidium (45–50; Madagascar, the Comoro Islands; in Pyrostria?), Pygmaeothamnus chamaedendrum (southern Africa), Canthium (c 50; tropical Africa, Madagascar, India, Sri Lanka), Rytigynia (125–130; tropical and and southern Africa), Cuviera (c 20; tropical West and Central Africa), Globulostylis (8; Central Africa), Vangueriella (18; tropical West and Central Africa), Pygmaeothamnus zeyheri (tropical and southern Africa), Multidentia (11; tropical Africa), Robynsia (1; R. glabrata; tropical West Africa), Vangueriopsis (4; tropical Africa), ’Vangueria’ (c 60; tropical Africa, Madagascar; non-monophyletic). – Unplaced Vanguerieae Meyna (11; eastern tropical Africa, the Comoro Islands and eastwards to Southeast Asia), Ancylanthos (1; A. rubiginosus; tropical Africa), Eriosemopsis (1; E. subanisophylla; KwaZulu-Natal, borner of Eastern Cape), Hutchinsonia (2; tropical West Africa), Perakanthus (1; P. velutinus; the Malay Peninsula), Cyclophyllum (c 40; Malesia and eastwards to tropical Australia, New Caledonia and Fiji). – Tropical regions in the Old World eastwards to tropical Australia, Melanesia and Polynesia.

[Greeneeae+[Aleisanthieae+Ixoreae]]

Greeneeae Mouly, J. Florence et B. Bremer in A. Mouly et al. in Ann. Missouri Bot. Gard. 96: 155. Apr 2009

2/8–9. Greenea (7–8; Southeast Asia, West Malesia), Spathichlamys (1; S. oblonga; Burma). – Southeast Asia, West Malesia.

[Aleisanthieae+Ixoreae]

Aleisanthieae Mouly, J. Florence et B. Bremer in A. Mouly et al. in Ann. Missouri Bot. Gard. 96: 155. Apr 2009

3/10. Aleisanthia (2; the Malay Peninsula), Aleisanthiopsis (2; Borneo), Greeniopsis (6; the Philippines). – West Malesia. – Aleisanthieae are sister to Ixoreae.

Ixoreae Benth. et Hook. f., Gen. Plant. 2: 9, 22. 7-9 Apr 1873

1/c 560. Ixora (c 560; tropical regions on both hemispheres).

Coffeeae alliance”

The “Coffeeae alliance” comprises Airospermeae, Augusteae, Alberteae, the [Nematostylis+Razafimandimbisonia] clade, the [Bertiera+Coffeeae] clade, the Octotropideae and the [Sherbournieae+Cordiereae+Pavetteae+Gardenieae] clade.

[Airospermeae+[Augusteae+[Alberteae+[[Nematostylis+Razafimandimbisonia]+[[Bertiera+Coffeeae]+[Octotropideae+[Sherbournieae+Cordiereae+Pavetteae+Gardenieae]]]]]]]

Airospermeae Kainul. et B. Bremer in K. Kainulainen, S. G. Razafimandimbison et B. Bremer, Bot. J. Linn. Soc. 173: 395. [DOI: 10.1111/boj.12038]. 25 Apr 2013.

2/7. Airosperma (6; New Guinea, Fiji), Boholia (1; B. nematostylis; Sumatra, the Philippines, Flores). – Malesia to New Guinea, Fiji. – This clade is sister to the remaining “Coffeeae alliance”.

[Augusteae+[Alberteae+[[Nematostylis+Razafimandimbisonia]+[[Bertiera+Coffeeae]+[Octotropideae+[Sherbournieae+Cordiereae+Pavetteae+Gardenieae]]]]]]

Augusteae Kainul. et B. Bremer in K. Kainulainen, S. G. Razafimandimbison et B. Bremer, Bot. J. Linn. Soc. 173: 395. [DOI: 10.1111/boj.12038]. 25 Apr 2013

2/c 85. Augusta (4; New Caledonia, Fiji, Mexico, northeastern and central Brazil), Wendlandia (c 80; northeastern Africa, Iraq to Malesia, tropical Australia). – Tropical and subtropical South Asia eastwards to Melanesia, tropical Australia, Mexico, Brazil. – This clade is sister to the remaining “Coffeeae alliance”.

[Alberteae+[[Nematostylis+Razafimandimbisonia]+[[Bertiera+Coffeeae]+[Octotropideae+[Sherbournieae+Cordiereae+Pavetteae+Gardenieae]]]]]

Alberteae Sond. in W. H. Harvey et O. W. Sonder, Fl. Cap. 3: 1. 24 Feb-30 Jun 1865

1/1. Alberta (1; A. magna; southern Africa). – Alberta is sister to the remaining “Coffeeae alliance”.

[[Nematostylis+Razafimandimbisonia]+[[Bertiera+Coffeeae]+[Octotropideae+[Sherbournieae+Cordiereae+Pavetteae+Gardenieae]]]]

[Nematostylis+Razafimandimbisonia]

2/6. Nematostylis (1; N. anthophylla; Madagascar), Razafimandimbisonia (5; Madagascar). – This clade is sister to the remaining “Coffeeae alliance”.

[[Bertiera+Coffeeae]+[Octotropideae+[Sherbournieae+Cordiereae+Pavetteae+Gardenieae]]]

[Bertiera clade+Coffeeae]

This clade is sister-group to the remaining “Coffeeae alliance”.

Bertiereae Bridson in Fl. Zambes. 5(3): 386. 2003

c 1/c 55? Bertiera pro parte (c 55?; tropical Africa, Madagascar, the Mascarene Islands, tropical America). – The Bertiera clade is sister to Coffeeae.

Coffeeae DC. in Ann. Mus. Natl. Hist. Nat. 9: 217. 1807 [‘Coffeaceae’]

11/c 280. Tricalysia (c 100; tropical Africa, Madagascar), Coffea (c 110; tropical Africa, Madagascar, the Mascarene Islands), Argocoffeopsis (8; tropical Africa), Calycosiphonia (2; tropical Africa), Belonophora (6; tropical Africa), Sericanthe (17; tropical Africa), Diplospora (≥10; tropical Asia), Pubistylus (1; P. andamanensis; the Andaman Islands), Discospermum (7; tropical Asia), Xantonnea (2–3; Southeast Asia), Nostolachma (c 10; tropical Asia). – Tropical Africa, Madagascar, the Mascarene Islands, the Andaman Islands, tropical Asia.

[[Burchellia+Galiniera]+[Mantalania+[Didymosalpinx+Monosalpinx]+[Octotropideae+[Sherbournieae+Cordiereae+Pavetteae+Gardenieae]]]]

[Burchellia+Galiniera]

Burchellia (1; B. bubalina; South Africa, Swaziland), Galiniera (2; tropical Africa, Madagascar). – This clade is sister-group to the remaining “Coffeeae alliance”.


[Mantalania+[Didymosalpinx+Monosalpinx]+[Octotropideae+[Sherbournieae+Cordiereae+Pavetteae+Gardenieae]]]

Mantalania clade

Mantalania (2–3; Madagascar), Pseudomantalania (1; P. macrophylla; Madagascar)?


[Didymosalpinx+Monosalpinx]

Monosalpinx (1; M. guillaumetii; tropical West Africa), Didymosalpinx (3; tropical Africa).


[Octotropideae+[Sherbournieae+Cordiereae+Pavetteae+Gardenieae]]

Octotropideae Bedd., Fl. Sylv. S. India 28: cxxvi, cxxxiv-12. 1874

c 28/c 125. Cremaspora (3–4; tropical Africa, the Comoro Islands); Feretia (2; tropical Africa), Kraussia (4; tropical and southern Africa), Octotropis (2; southern India, Burma), Xantonneopsis (1; X. robinsonii; Southeast Asia), Petitiocodon (1; P. parviflorus; Nigeria, Cameroun)?, Canephora (5; Madagascar), Chapelieria (3; Madagascar), Fernelia (4; the Mascarene Islands), Flagenium (6; Madagascar), Nargedia (1; N. macrocarpa; Sri Lanka), Paragenipa (1; P. lancifolia; the Seychelles), Lamprothamnus (1; L. zanguebaricus; tropical East Africa), Jovetia (2; Madagascar), Lemyrea (4; Madagascar)?, Polysphaeria (c 20; tropical Africa, Madagascar, the Comoro Islands), Cowiea (2; Borneo, the Philippines), Gallienia (1; G. sclerophylla; Madagascar), Hypobathrum (c 35; tropical Asia), Hyptianthera (2; northern India, Thailand), Morindopsis (2; Southeast Asia), Pouchetia (6; tropical Africa), Ramosmania (2; Rodrigues), Rhadinopus (2; New Guinea), Scyphostachys (2; Sri Lanka), Villaria (6; the Philippines), Zuccarinia (1; Z. macrophylla; West Malesia). – Tropical Africa, Madagascar, the Mascarene Islands, the Seychelles, India, Sri Lanka, Southeast Asia, Malesia, with their highest diversity in tropical Africa and Madagascar. – Cremaspora is sister to the remaining Octotropideae. – This clade is sister-group to the clade [Sherbournieae+Cordiereae+Pavetteae+Gardenieae].

[Sherbournieae+Cordiereae+Pavetteae+Gardenieae]

Pavetteae Dumort., Anal. Fam. Plant.: 32. 1829

c 17/c 640. Nichallea (1; N. soyauxii; tropical Africa), Coleactina (1; C. papalis; Central Africa), Homollea (3; Madagascar), Homolliella (1; H. sericea; Madagascar), Dictyandra (2; tropical West and Central Africa), Leptactina (19; tropical and southern Africa), Pavetta (c 300; tropical regions in the Old World), Cladoceras (1; C. subcapitata; tropical East Africa), Rutidea (22; tropical Africa, Madagascar), Coptosperma (c 50; tropical Africa), Robbrechtia (2; Madagascar), Schizenterospermum (4; Madagascar), Paracephaelis (3; Madagascar), Tennantia (1; T. sennii; tropical East Africa), Tarenna (c 200; tropical regions in the Old World). – Unplaced Pavetteae Pachystylus (2; New Guinea)?

Gardenieae A. Rich. ex DC., Prodr. 4: 342, 367. late Sep 1830 [‘Gardeniaceae’]

50–55/520–530. Schumanniophyton (3; tropical Africa)?; Ceriscoides (11; tropical Asia), Genipa (3; tropical America), Brenania (2; Central Africa), ’Gardenia’ (c 200; tropical and subtropical regions in the Old World eastwards to Pacific islands; non-monophyletic), Larsenaikia (3; Northern Territory, northern and eastern Queensland), Kailarsenia (c 10; tropical Asia, eastern Queensland), Coddia (1; C. rudis; South Africa, Swaziland), Aulacocalyx (9; tropical Africa), Heinsenia (1; H. diervilleoides; tropical Africa), Rothmannia (c 30; tropical and southern Africa and eastwards to the Seychelles and the Malay Peninsula), Phellocalyx (1; P. vollessenii; Tanzania, Malawi, Mozambique), Kochummenia (2; the Malay Peninsula), Morelia (1; M. senegalensis; tropical Africa), Hyperacanthus (4; southeastern Africa, Madagascar), Aidia (c 50; tropical regions in the Old World), Aoranthe (5; tropical Africa), Benkara (c 32; tropical and subtropical Asia), Massularia (1; M. acuminata; tropical Africa), Tocoyena (12; tropical America), Sphinctanthus (8; tropical South America), Rosenbergiodendron (3; tropical America), ‘Casasia’ (11; Central America, the West Indies; non-monophyletic), ’Randia’ (c 20; tropical and subtropical America from Florida and southwards to Bolivia; non-monophyletic), Melanoxerus (1–2; Madagascar), Preussiodora (1; P. sulphurea; tropical West Africa), Pleiocoryne (1; P. fernandensis; tropical West and Central Africa), Oligocodon (1; O. cunliffeae; tropical West and Central Africa), Euclinia (2; tropical Africa), Macrosphyra (3; tropical Africa), Calochone (2; Central Africa), Brachytome (4; southern China, tropical Asia), Dioecrescis (1; D. erythroclada; India), Tamilnadia (1; T. uliginosa; South and Southeast Asia), Vidalasia (5; Southeast Asia, the Philippines), Rubovietnamia (2; Guangxi, Vietnam), Duperrea (2; India, southern China, Southeast Asia), Porterandia (5–6; West Malesia), Tarennoidea (2; tropical Asia), Atractocarpus (c 25; the Philippines, New Caledonia), Deccania (1; D. pubescens; India), Catunaregam (8; tropical Africa, tropical Asia), Bungarimba (4; Malaysia, Sumatra, Borneo, New Guinea), Trukia (6; Micronesia, central Polynesia), Sukunia (1; S. longipes; Fiji). – Unplaced Gardenieae Adenorandia (1; A. kalbreyeri; tropical Africa)?, Aidiopsis (2; Malesia)?, Alleizettella (2; southeastern China, northern Vietnam)?, Fosbergia (5; Burma, Yunnan, Thailand, Vietnam)?, Ganguelia (1; G. gossweileri; Angola)?, Gardeniopsis (1; G. longifolia; West Malesia)?, Himalrandia (3; Himalayas)?, Pseudaidia (1; P. speciosa; India)? – Pantropical.

Sherbournieae Mouly et B. Bremer in Taxon 63: 814. 28 Aug 2014

4/c 60. Atractogyne (3; tropical West and Central Africa), Mitriostigma (5; tropical and southern Africa), Oxyanthus (c 40; tropical and subtropical Africa), Sherbournia (c 10; tropical Africa). – Seeds with folded testa.


Cordiereae A. Rich. ex DC., Prodr. 4: 342, 445. Late Sep 1830

12/c 135. Botryarrhena (2; Venezuela, Brazil, Peru), Stachyarrhena (13; tropical America), Stenosepala (1; S. hirsuta; Panamá, Colombia), Kutchubaea (11; tropical South America), Glossostipula (2; southern Mexico, Guatemala), Duroia (c 25; tropical America), Amaioua (c 25; tropical South America), Riodocea (1; R. pulcherrima; Brazil), Agouticarpa (7; Costa Rica to Bolivia), Melanopsidium (1; M. nigrum; Brazil), Cordiera (9; northern South America), Alibertia (c 35; tropical America) – Unplaced Cordiereae Borojoa (c 10; tropical America)? – Tropical America.

Rubioideae Bremek. ex Verdc. in Bull. Jard. Bot. État Bruxelles 28: 280. 30 Sep 1958

174–181/6.200–6.400. Cosmopolitan. Usually herbs. Root phellogen usually deeply seated (in e.g. Paederia superficial). Raphides present. Hairs septate. Corolla with valvate aestivation. Heterostyly frequently occurring. Pollen grains usually bicellular (sometimes tricellular) at dispersal. Integument one to 14 cell layers thick (rarely absent). Antipodal cells usually not persistent. Suspensor haustorium sometimes present. The atpB promoter absent (lost). Shikimic acid-derived anthraquinones and cyclotide proteins present. Route II carboxylated iridoids and indole alkaloids absent. Aluminium often accumulated (particularly in woody species).

Colletoecemateae Rydin et B. Bremer in C. Rydin et al. in Plant Syst. Evol. 278: 120. Mar 2009

1/1. Colletoecema (1; C. dewevrei; Central Africa). – Colletoecema is sister to the remaining Rubioideae, according to, e.g., Robbrecht & Manen (2006) and Rydin & al. (2008).

Ophiorrhizeae Bremek. ex Verdc. in Bull. Jard. Bot. État Bruxelles 28: 281. 30 Sep 1958

7–8/210–220. Lerchea (8; Sumatra, Java), Xanthophytum (32; Java, Borneo and eastwards to Fiji), Kajewskiella (2; the Solomon Islands), Keenania (5; Assam, Southeast Asia; incl. Myrioneuron?), Myrioneuron (9–14; northern India, Nepal, Bhutan, southern China, Vietnam; in Keenania?), Neurocalyx (5; Sri Lanka, southern India), Ophiorrhiza (c 150; tropical Asia). – Tropical Asia and eastwards to Melanesia. – Ophiorrhiza has the atpB promotor region which is absent from other Rubioideae.

Urophylleae Bremek. ex Verdc. in Bull. Jard. Bot. État Bruxelles 28: 281. 30 Sep 1958

c 14?/190–250. Temnopteryx (1; T. sericea; Central Africa), Raritebe (1; R. palicoureoides; tropical America), Amphidasya (7; tropical America), ’Urophyllum’ (85–150; tropical regions in the Old World and northwards to China and Japan; polyphyletic), Pentaloncha (3; tropical West and Central Africa), Pauridiantha (c 30; tropical Africa), Praravinia (c 50; Borneo, the Philippines, Sulawesi), Antherostele (4; the Philippines)?, Crobylanthe (1; C. pellacalyx; Borneo)?, Didymopogon (1; D. sumatranum; Sumatra)?, Lepidostoma (1; L. polythyrsum; Sumatra)?, Rhaphidura (1; R. lowii; Borneo)?, Rhipidantha (1; R. chlorantha; Uluguru Mountains in Tanzania)?, Stichianthus (1; S. minutiflorus; Borneo)? – Temnopteryx may be sister to the remaining Urophylleae, although seven of the genera in Urophylleae are not yet analyzed. Raritebe is sister to Amphidasya and Pentaloncha sister to Pauridiantha.

[[Lasiantheae+Perama]+[Coussareeae+[”Spermacoceae alliance”+”Psychotrieae alliance”]]]

[Lasiantheae+Perama]

Lasiantheae B. Bremer et Manen in Plant Syst. Evol. 225: 57. 10 Jan 2001

c 4/c 210. Lasianthus (c 170; tropical Africa, tropical Asia and eastwards to tropical Australia), Trichostachys (14; tropical Africa), 'Ronabea' (3; Central America, tropical South America to Bolivia), Saldinia (22; Madagascar). – Tropical regions in the Old World.

Perameae Bremek. ex S. P. Darwin in Taxon 25: 605. 26 Nov 1976

1/9–14. Perama (9–14; tropical America). – Perama is sister to Lasiantheae.

[Coussareeae+[”Spermacoceae alliance”+”Psychotrieae alliance”]]

Coussareeae Benth. et Hook. f., Gen. Plant. 2: 9, 24. 7-9 Apr 1873

7–8/305–310. Coccocypselum (c 20; tropical America), Declieuxia (27; tropical America), Hindsia (11; Brazil), Coussarea (c 110; tropical America), Faramea (c 130; tropical America), Cruckshanksia (7; northern Chile, northern Argentina; incl. Oreopolus?), Oreopolus (1; O. glacialis; the Andes; in Cruckshanksia?), Heterophyllaea (8; Bolivia, Argentina). – Tropical America.

The “Spermacoceae alliance”

[Danaideae+[[Knoxieae+Spermacoceae]+[Anthospermeae+[Argostemmateae+[Dunnieae+[Paederieae+[Plocama+[Theligonum+Rubieae]]]]]]]]

Danaideae B. Bremer et Manen in Plant Syst. Evol. 225: 60. 10 Jan 2001

3/c 70. Danais (c 40; Tanzania, Madagascar, the Mascarene Islands), Schismatoclada (c 20; Madagascar), Payera (10; Madagascar). – Tanzania, Madagascar, the Mascarene Islands, with their highest diversity on Madagascar.

[[Knoxieae+Spermacoceae]+[Anthospermeae+[Argostemmateae+[Dunnieae+[Paederieae+[Plocama+[Theligonum+Rubieae]]]]]]]

[Knoxieae+Spermacoceae]

Knoxieae Benth. et Hook. f., Gen. Plant. 2: 9, 21. 7-9 Apr 1873

c 14/130–140. Chamaepentas (1; C. greenwayi; tropical East Africa), ’Carphalea’ (15; tropical Africa, Madagascar, Socotra; paraphyletic), Triainolepis (2; tropical East Africa, Madagascar), Parapentas (4–5; tropical Africa, Madagascar?), Dolichopentas (4; tropical Africa), Phyllopentas (13; tropical Africa, Madagascar), Knoxia (9; tropical Africa, tropical Asia), Pentas (c 35; tropical Africa, Madagascar, the Arabian Peninsula), Rhodopentas (2; tropical East Africa), Chlorochorion (2; tropical Africa), Pentanisia (15–20; tropical Africa, Madagascar), Batopedina (3; western and southern tropical Africa), Otomeria (8; tropical Africa, Madagascar), Otiophora (c 20; tropical Africa, Madagascar). – Knoxieae are sister to Spermacoceae.

Spermacoceae Cham. et Schltdl. ex DC., Prodr. 4: 343, 538. late Sep 1830

60–65/c 1.150. Kohautia (27; tropical and subtropical Africa, the Arabic Peninsula to India, Thailand, tropical Australia), Lathraeocarpa (2; southern Madagascar), Phylohydrax (2; tropical East Africa, Madagascar), Conostomium (4; tropical East Africa), Amphiasma (5–6; tropical and southwestern Africa), Pentanopsis (2; northeastern tropical Africa; in Oldenlandia?), Dentella (8; tropical Asia to tropical Australia and New Caledonia), Pentodon (2; tropical and subtropical regions in Africa, the Seychelles, the Arabian Peninsula, tropical and subtropical America), Edrastima (5; tropical and subtropical Africa, tropical Asia, North and South America), Hedyotis (c 180; tropical and subtropical Asia eastwards to New Guinea, island in the northwestern Pacific), Dibrachionostylus (1; D. kaessneri; tropical East Africa), Neanotis (c 30; tropical Asia to tropical Australia, islands in the Pacific), Involucrella (2; Southeast Asia), Debia (4; Southeast Asia, the Andaman Islands, the Philippine Islands), Kadua (c 30; French Polynesia, the Hawaiian Islands), Leptopetalum (8; tropical Asia to tropical Australia, islands in the Pacific), Scleromitrion (?; tropical Asia to tropical Australia, islands in the Pacific), Exallage (15; tropical Asia to tropical Australia, islands in the Pacific), Dimetia (7; tropical Asia), Cordylostigma (9; tropical Africa, Madagascar), ‘Oldenlandia’ (c 300?; tropical and subtropical regions on both hemispheres; polyphyletic) Phialiphora (2; northwestern Madagascar), Cordylostigma (9; tropical Africa, Madagascar), Stenaria (5; North America), Thecorchus (1; T. wauensis; tropical Africa), Agathisanthemum (5–6; tropical Africa, the Comoro Islands), Dentella (10; tropical Asia and eastwards to tropical Australia and New Caledonia), Pentodon (2; tropical and subtropical regions in Africa, the Seychelles, the Arabian Peninsula, tropical and subtropical America), Kohautia (c 60; tropical regions in the Old World), Lathraeocarpa (2; southern Madagascar), Phylohydrax (2; tropical East Africa, Madagascar), Gomphocalyx (1; G. hernarioides; Madagascar), Manostachya (3; tropical Africa), Conostomium (4; tropical East Africa), Pentanopsis (2; northeastern tropical Africa), Amphiasma (5–6; tropical and southwestern Africa), Hedythyrsus (2; tropical Africa), Pseudonesohedyotis (1; P. bremekampii; Uuguru Mountains in Tanzania), Mitrasacmopsis (1; M. quadrivalvis; tropical East Africa, Madagascar), Astiella (1; A. delicatula; Madagascar), Synaptantha (2; tropical and subtropical regions in Australia), Dibrachionostylus (1; D. kaessneri; tropical East Africa), Arcytophyllum (17; southern Mexico to Bolivia), Houstonia (20; North America), Mitracarpus (c 30; tropical America), Richardia (c 15; tropical America), ‘Spermacoce’ (c 250; tropical and subtropical regions on both hemispheres; paraphyletic), Hydrophylax (1–3; H. maritima; tropical and subtropical Africa, Madagascar, India, Thailand), Bouvardia (c 20; tropical America), Diodia (c 30; tropical and subtropical regions in Africa and America), Galianthe (50–55; tropical Central and South America), Ernodea (9; southeastern United States, the West Indies; in Spermacoce?), Mitracarpus (c 30; tropical America; in Spermacoce?), Psyllocarpus (9; Brazil; in Spermacoce?), Manettia (c 80; tropical America), Nesohedyotis (1; N. arborea; St. Helena), Psyllocarpus (9; Brazil), Emmeorhiza (1; E. pohliana; tropical South America), Denscantia (4; Brazil), Anthospermopsis (1; A. catechosperma; Brazil), Staelia (16; northern South America), Pleiocraterium (4; southern India, Sri Lanka, Sumatra), Neanotis (28; tropical Asia and eastwards to tropical Australia), Carterella (1; C. alexanderae; Baja California in northwestern Mexico), Dolichometra (1; D. leucantha; eastern Usambara Mountains in Tanzania), Leptoscela (1; L. ruellioides; eastern Brazil), Lucya (1; L. tetrandra; the West Indies), Phyllocrater (1; P. gibbsiae; Borneo), Polyura (1; P. geminata; Assam), Stephanococcus (1; S. crepinianus; tropical Africa), Oldenlandiopsis (1; O. callitrichoides; the West Indies), Micrasepalum (2; Cuba, Hispaniola), Spiradiclis (38; India, southwestern China, Southeast Asia, Java), Crusea (13; southwestern United States, Mexico, Central America), Leptomischus (3–7; Southeast Asia), Merumea (2; Guayana Highlands). – Tropical and subtropical regions on both hemispheres, North America.

[Anthospermeae+[Argostemmateae+[Dunnieae+[Paederieae+[Plocama+[Theligonum+Rubieae]]]]]]

Anthospermeae Cham. et Schltdl. in Linnaea 3: 309. Oct 1828

10/c 190. Anthospermum (c 40; Africa, Madagascar), Nenax (11; southern Namibia, South Africa), Galopina (4; southern Africa to Malawi), Phyllis (2; Madeira, the Canary Islands), Coprosma (c 100; Kerguelen Island, southern China, southeastern South Australia, eastern New South Wales, Victoria, Tasmania, New Zealand, Stewart Island, Macquarie Island, Campbell Island, Auckland Islands, Antipode Islands, the Hawaii Islands, southern South America, Tristan da Cunha), Durringtonia (1; D. paludosa; southeastern Queensland, northeastern New South Wales), Pomax (1; P. umbellata; Australia), Leptostigma (6; Southeast Asia, New Zealand, western South America), Carpacoce (7; Western and Eastern Cape), Opercularia (c 18; southwestern Western Australia, southeastern South Australia to southeastern Queensland, Tasmania). – Macaronesia, Africa, Madagascar, Southeast Asia, Australia, Tasmania, New Zealand, the Hawaiian Islands, western and southern South America, Subantarctic islands.

[Argostemmateae+[Dunnieae+[Paederieae+[Plocama+[Theligonum+Rubieae]]]]]

Argostemmateae Bremek. ex Verdc. in Bull. Jard. Bot. État Bruxelles 28: 281. 30 Sep 1958

3/c 130. Mouretia (4; Southeast Asia), Mycetia (c 25; tropical Asia), Argostemma (c 100; tropical regions in the Old World). – Tropical regions in the Old World. Anthers in Argostemma connate.

[Dunnieae+[Paederieae+[Plocama+[Theligonum+Rubieae]]]]

Dunnieae Rydin et B. Bremer in C. Rydin et al. in Plant Syst. Evol. 278: 121. Mar 2009

2/5. Dunnia (4; northern India, southeastern China), Foonchewia (1; F. guangdongensis; Guangdong). – Northern India, southern China.

[Paederieae+[Plocama+[Theligonum+Rubieae]]]

Paederieae DC., Prodr. 4: 342, 470. late Sep 1830

6/110–115. Saprosma (45–50; tropical Asia), Paederia (c 30; tropical regions on both hemispheres), Spermadictyon (1; S. suaveolens; India), Leptodermis (c 30; Himalayas to Japan), Serissa (2; southern China), Pseudopyxis (2; Japan). – Tropical regions on both hemispheres.

[Plocama+[Theligonum+Rubieae]]

Putorieae Lange in H. M. Willkomm et J. M. C. Lange, Prodr. Fl. Hispan. 2: 299. 1870

1/c 35. Plocama (c 35; the Canary Islands, the Mediterranean, southwestern Asia eastwards to northwestern India). – Plocama is sister to [Theligonum+Rubieae].

[Theligoneae+Rubieae]

Theligoneae Baill., Hist. Plant. 4: 46, 55. 1872

1/3–4. Theligonum (3–4; Macaronesia, the Mediterranean, southwestern China, Japan). – Usually unisexual (monoecious), annual or perennial herbs. Calyx strongly reduced. Corolla bilobate to quinquelobate. Stamens (two to) seven to twelve (to 30), sometimes basally fused in fascicles of two, four or six. Pollen grains 4–8-zonoporate. Pistil composed of two connate carpels (one carpel fertile, the second carpel aborted: pseudomonomery). Style gynobasic. Placentation basal. Ovule solitary, campylotropous or amphitropous. Fruit a nutlike drupe. Pericarp with elaiosome. Endosperm starchy. Iridoids present. – Theligonum is sister to Rubieae.

Rubieae Baill., Hist. Plant. 7: 365, 390. Feb 1880

4?/c 700. Kelloggia (2; K. chinensis: southern China, Bhutan; K. galioides: southwestern United States, northwestern Mexico); Didymaea (5; Central America), Rubia (60–80; Macaronesia, the Mediterranean, Africa, temperate Asia, North America), Galium (>600; cosmopolitan). – Cosmopolitan. – Rubieae have usually foliaceous stipules resembling verticillate leaves. Kelloggia may be sister to the remaining Rubieae. Didymaea, Galium (sensu lato) and Rubia form a polytomy in the analysis by Ehrendorfer & Barfuss (2014).

The “Psychotrieae alliance”

[Craterispermeae+Schradereae+[Gaertnereae+[Mitchelleae+Morindeae]]+Prismatomerideae+Psychotrieae]

Craterispermeae Verdc. in Bull. Jard. Bot. Bruxelles 28: 281. 30 Sep 1958

1/16. Craterispermum (16; tropical Africa, Madagascar, the Seychelles).

Schradereae Bremek. in Rec. Trav. Bot. Néerl. 31: 253. 1934

3/c 60. Schradera (c 55; Malesia, tropical America), Lecananthus (3; West Malesia), Leucocodon (1; L. reticulatum; Sri Lanka). – Sri Lanka, Malesia, tropical America.

[Gaertnereae+[Mitchelleae+Morindeae]]

Gaertnereae Endl., Gen. Plant.: 577. Aug 1838

2/c 95. Pagamea (c 25; tropical South America), Gaertnera (c 70; tropical regions in the Old World). – Pantropical. – Gaertnereae are sister-group to the clade [Mitchelleae+Morindeae].

[Mitchelleae+Morindeae]

Mitchelleae Razafim. et B. Bremer in S. G. Razafimandimbison, C. Rydin et B. Bremer, Mol. Phylogen. Evol. 48: 221. Jul 2008

2/11–13. Damnacanthus (8–10; China, Japan, Taiwan), Mitchella (3; M. undulata: southern Korea, Japan, Taiwan; M. repens: North America; M. ovata: Ecuador). – East Asia, North America, Ecuador. – Mitchelleae are sister to Morindeae.

Morindeae Burnett, Outlines Bot.: 906. Feb 1835

c 6/c 170. Appunia (6–8; southern Mexico, Central America, the West Indies, tropical South America), ‘Morinda’ (c 130; tropical and subtropical regions on both hemispheres; polyphyletic), Gynochthodes (>20; Southeast Asia, Malesia and eastwards to tropical Australia and Melanesia; incl. Morinda pro parte), Coelopyrena (1; C. salicifolia; East Malesia), Coelospermum (9; Southeast Asia, Malesia and eastwards to eastern Queensland and New Caledonia), Siphonandrium (1; S. intricatum; New Guinea). – Tropical and subtropical regions on both hemispheres.

Prismatomerideae Y. Z. Ruan in Acta Phytotaxon. Sin. 26: 446. Dec 1988 [‘Prismatomereae’]

4/22. Prismatomeris (16; Sri Lanka, Assam, Southeast Asia, Hainan, West Malesia), Gentingia (1; G. subsessilis; northwestern Malay Peninsula), Motleyia (1; M. borneensis; northwestern Borneo), Rennellia (4–5; the Malay Peninsula, Sumatra, Borneo). – Sri Lanka, Assam, Southeast Asia, Hainan, West Malesia.

Psychotrieae Cham. et Schltdl. in Linnaea 4: 4. Jan 1829 [‘Psychotriaceae’]

25–30/2.400–2.500. Schizocolea (2; tropical West Africa); ’Psychotria’ (1.900–2.000; tropical regions on both hemispheres; non-monophyletic), Gillespiea (1; G. speciosa; Fiji; in Psychotria?), Hedstromia (1; H. latifolia; Fiji; in Psychotria?), Notopleura (c 100; tropical America; in Psychotria?), Cremocarpon (9; Madagascar, the Comoro Islands), Pyragra (2; Madagascar), Amaracarpus (22; the Seychelles, tropical Asia and eastwards to tropical Australia and the Solomon Islands), Dolianthus (13; mountains in Papua New Guinea), Myrmecodia (c 25; Malesia and eastwards to Fiji), Hydnophytum (55; tropical Asia, with their highest diversity on New Guinea), Squamellaria (3–4; Fiji), Myrmephytum (8; the Philippines, Sulawesi, western New Guinea), Anthorrhiza (9; New Guinea), Calycosia (5; Polynesia), Palicourea (>200; tropical America), Rudgea (c 70; tropical America), Carapichea (2; tropical America), Margaritopsis (c 50; tropical regions on both hemispheres), Hymenocoleus (12; tropical Africa), ‘Geophila’ (c 20; tropical regions on both hemispheres; polyphyletic), ‘Chassalia’ (40–45; tropical regions in the Old World, with their largest diversity on Madagascar; polyphyletic), Coccochondra (1; C. laevis; Guayana Highlands), Peripeplus (1; P. bracteosus; Central Africa), Streblosa (c 25; West Malesia). – Pantropical. Anthorrhiza, Hydnophytum, Myrmecodia, Myrmephytum and Squamellaria are myrmecophilous with ant colonies living in the hollow stems. – Schizocolea is sister to the remaining Psychotrieae.

Unplaced Rubiaceae Acrobotrys (1; A. discolor; Colombia), Aphanocarpus (1; A. steyermarkii; Venezuela), Benzonia (1; B. corymbosa; Guinea), Berghesia (1; B. coccinea; Mexico), Bradea (5; Brazil), Byrsophyllum (2; India, Sri Lanka), Clarkella (2; Himalayas, Thailand), Coptophyllum (14; India, West Malesia), Coryphothamnus (1; C. auyantepuiensis; southeastern Venezuela), Cyanoneuron (5; Borneo, Sulawesi), Diacrodon (1; D. compressus; Brazil), Didymochlamys (2; Panamá, Colombia, Venezuela), Duidania (1; D. montana; Venezuela), Eizia (1; E. mexicana; Mexico), Etericius (1; Guyana; nomen dubium), Fergusonia (1; F. tetracocca; southern India, Sri Lanka), Flexanthera (2; Colombia, Bolivia), Holstianthus (1; H. barbigularis; Guayana Highlands), Klossia (1; K. montana; the Malay Peninsula), Lecariocalyx (1; L. borneensis; Borneo), Maschalodesme (2; New Guinea), Nodocarpaea (1; N. radicans; Cuba), Pagameopsis (2; Venezuela), Placocarpa (1; P. mexicana; Mexico), Pseudohamelia (1; P. hirsuta; the Andes), Riqueuria (1; R. avenia; Peru), Sacosperma (2; tropical Africa), Schwendenera (1; S. tetrapyxis; southeastern Brazil), Stachyococcus (1; S. adinanthus; Peru), Standleya (4; Brazil), Streblosiopsis (1; S. cupulata; Borneo), Stylosiphonia (2; Central America), Temnocalyx (1; T. nodulosa; Tanzania), Tobagoa (1; T. maleolens; Panamá, Tobago, Venezuela), Tortuella (1; T. abietifolia; Île Tortue near Hispaniola).

Cladogram (simplified) of Rubiaceae based on DNA sequence data (Bremer & Eriksson 2009, with Prismatomerideae added). ’Gardenieae’ are non-monophyletic. According to more recent analyses covering a larger part of the Rubiaceae taxa, Condamineeae are sister to the remaining Ixoroideae, and Mussaendeae and Sabiceeae are sister-groups.


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