RUBIACEAE 茜草科 qian cao ke Chen Tao (陈涛)1, Zhu Hua (朱华)2, Chen Jiarui (陈家瑞 Chen Chia-jui)3; Charlotte M. Taylor4, Friedrich Ehrendorfer5, Henrik Lantz6, A. Michele Funston4, Christian Puff5 Trees, shrubs, annual or perennial herbs, subshrubs, vines, or lianas, infrequently monocaulous or creeping and rooting at nodes, terrestrial or infrequently epiphytic, with bisexual flowers, infrequently dioecious, or rarely polygamo-dioecious (Diplospora, Galium, Guettarda, perhaps Brachytome) or monoecious (Galium), evergreen or sometimes deciduous (Hymenodictyon), sometimes armed with straight to curved spines (formed by modified stems or peduncles), infrequently with elongated principal stems bearing lateral short shoots (i.e., brachyblasts; Benkara, Catunaregam, Ceriscoides, Himalrandia, Leptodermis, Serissa), infrequently with lateral branches or short shoots spinescent (i.e., prolonged, sharp, and leafless at apex), infrequently with reduced internodes that give an appearance of verticillate leaf arrangement (Brachytome, Damnacanthus, Duperrea, Rothmannia, Rubovietnamia), infrequently with buds resinous (Gardenia) or mucilaginous (Scyphiphora), infrequently with tissues fetid when bruised, [rarely with swollen hollow stems or leaf bases housing ants (Neonauclea)]; branchlets terete to angled or quadrate, in latter two cases often becoming terete with age, or rarely flattened (Wendlandia) or winged (Hedyotis, Rubia), buds conical or rounded with stipules valvate or imbri- cate, or infrequently flattened with stipules erect and pressed together (Cinchona, Haldina, Nauclea, Neonauclea). Raphides present or absent. Leaves opposite, verticillate, or apparently verticillate (i.e., closely set due to reduced internodes), decussate or occasion- ally distichous, petiolate to sessile, infrequently somewhat to strongly anisophyllous, rarely punctate- or striate-glandular (Galium); margins flat to occasionally undulate or crisped, entire or rarely lobed (Hymenodictyon, Morinda) to denticulate or serrate (Hymeno- dictyon, Leptomischus, Ophiorrhiza, Wendlandia); secondary veins pinnate or rarely triplinerved or palmate (Hedyotis, Rubia), free (i.e., eucamptodromous) or uniting near margins (i.e., brochidodromous) in weak to well-developed or rarely substraight submarginal vein, sometimes with foveolate (i.e., pitted or cryptlike) and/or tufted (i.e., pubescent) domatia (i.e., structures that house mites) in abaxial axils, these rarely also present in axils of tertiary veins (Morinda), with presence of domatia often variable within a species; tertiary and/or quaternary venation rarely arranged in regular squares (Guettarda), regular rectangles (i.e., clathrate; Urophyllum), or lineolate (i.e., closely parallel within each areole; Timonius); petiole rarely articulate at base (Ixora); stipules persistent with leaves, deciduous before leaves, or quickly caducous, interpetiolar and infrequently fused to adjacent petioles or leaf bases, sometimes united around stem into a sheath, rarely completely united into a conical cap (i.e., calyptrate; Gardenia), with interpetiolar portion variously triangular in general shape to truncate, with apex entire or bilobed, multifid, lacerate, setose, or laterally appendaged, with apex, lobes, setae, and/or appendages sometimes glandular (Chassalia, Hedyotis, Hymenodictyon, Knoxia, Mitchella, Mycetia, Neanotis, Ophiorrhiza, Pentas, Pseudopyxis, Psychotria, Trailliaedoxa), internally (i.e., adaxially) with small to well-developed col- leters (i.e., glandular trichomes), these infrequently persistent after stipules fall (Psychotria), or stipules rarely expanded into 1 to several leaflike segments and then apparently absent due to leaflike form that gives an appearance of verticillate leaves (Argostemma, Asperula, Galium, Microphysa, Phuopsis, Rubia). Inflorescences terminal, axillary (i.e., borne at both axils at a node), or pseudo- axillary (i.e., borne consistently in 1 axil per node; lateral), sometimes apparently leaf-opposed due to marked anisophylly, or rarely superaxillary (Damnacanthus, Diplospora) or cauline (Mycetia), variously cymose to thyrsiform, corymbiform, paniculiform, racemiform, spiciform, fasciculate, or capitate and few to many flowered or occasionally reduced to a solitary flower, pedunculate (peduncle here used for basalmost axis supporting inflorescence or solitary flower) to sessile, when sessile often with 3 principal axes (i.e., tripartite), bracteate or bracts sometimes reduced or apparently absent, with bracts (here usually including bracts borne on pedi- cels or next to flowers, i.e., bracteoles) generally triangular to linear or sometimes leaflike (i.e., similar to normal or somewhat reduced leaves) and rarely enlarged, petaloid, and resembling calycophylls (Dunnia, Neohymenopogon), infrequently fused and invo- lucral, occasionally fused in pairs (i.e., forming a calyculus, or calyculate), infrequently thickened and spatulate to clavate or conical (genera of Naucleeae), or infrequently stipuliform, rarely glandular (Damnacanthus, Mycetia), multifid to fimbriate (Damnacanthus, Kelloggia, Spermacoce) or spinescent (Phuopsis). Flowers sessile to pedicellate (pedicel here used for ultimate axis immediately supporting a single flower, except when this is a peduncle), bisexual and monomorphic, distylous, or rarely tristylous (Chassalia, Pentas), unisexual with 2 forms generally similar except for corolla size and hypanthium development, or rarely cleistogamous (Ophiorrhiza), actinomorphic or rarely zygomorphic (Argostemma), sometimes with ovaries of individual flowers partially to fully fused (Mitchella, Morinda, Mouretia, Nauclea), variously diurnal or nocturnal, usually sweetly fragrant, protandrous [or rarely proto- gynous], occasionally with secondary pollen presentation (e.g., Ixora, Pavetta, Phuopsis, Scyphiphora, genera of Naucleeae). Calyx gamosepalous and fused to inferior ovary in hypanthium or ovary portion, this generally ellipsoid, turbinate, obconic, cylindrical, or occasionally subglobose to hemispherical, glabrous and smooth, pubescent, or rarely tuberculate (Galium) or with unusual flattened (Dentella) or hooked, sometimes glandular trichomes (Galium, Kelloggia), or rarely longitudinally ridged to winged (Gardenia, 1 Fairy Lake Botanical Garden, 160 Xianhu Road, Liantang, Shenzhen, Guangdong 518004, People’s Republic of China. 2 Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Menglun Town, Mengla County, Xishuangbanna, Yunnan 666303, People’s Republic of China. 3 Institute of Botany, Chinese Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’s Republic of China. 4 Missouri Botanical Garden, P.O. Box 299, Saint Louis, Missouri 63166-0299, U.S.A. 5 Department of Systematic and Evolutionary Botany, Faculty of Life Sciences, University of Vienna, Rennweg 14, A-1030 Vienna, Austria. 6 Museum of Evolution, Botany Section (Fytoteket), Evolutionary Biology Center, Uppsala University, Norbyvägen 16, SE-752 36 Uppsala, Sweden. 57 58 RUBIACEAE Spiradiclis), above this developed into a free limb portion, this limb variously tubular to cupular or infrequently reduced to obsolete (Asperula, Coffea, Galium, Leptunis, Microphysa, Ophiorrhiza, Phuopsis, Rubia), truncate to 4- or 5(–9)-denticulate (i.e., lobes reduced to tiny projections along a generally truncate margin) or shallowly to deeply 4- or 5(–9)-lobed, open in bud or rarely with lobes markedly imbricate (Emmenopterys, Keenania), [or rarely spathaceous (i.e., fused into a conical cap that splits irregularly)], inside variously near base and/or at sinuses between lobes with few to numerous small colleters, rarely densely or markedly veined (Clarkella, Myrioneuron, Pseudopyxis), rarely on margins with well-developed, sessile to stalked glands (Mycetia), lobes generally triangular to linear, occasionally obtuse to lanceolate or oblanceolate, or rarely prolonged into a slender shaft bearing a thickened apical portion (Neonauclea), occasionally slightly to markedly unequal on an individual flower with all lobes of different lengths or infrequently in unequal pairs of similar lengths (Diodia, Mitracarpus, Spermacoce), infrequently with 1(to 5, Mussaenda) lobe on some (or all, Mussaenda) flowers of an inflorescence enlarged into a calycophyll (i.e., a membranous to papery, petaloid, veined, white to colored blade borne on a generally well-developed stipe; Emmenopterys, Morinda, Mussaenda, Schizomussaenda). Corolla large and often showy to reduced, gamopetalous, white, yellow, orange, red, blue, purple, and/or pale green, when nocturnal often white at anthesis becoming yellow with age, variously funnelform, salverform (i.e., hypocrateriform), tubular, campanulate, or occa- sionally rotate to infrequently urceolate (i.e., swollen in basal part of tube; Canthium, Lasianthus) or inflated (i.e., markedly swollen in middle or upper part of tube; Keenania, Leptomischus), infrequently curved in tube and/or gibbous (i.e., asymmetrically swollen at very base of tube; Chassalia, Guettarda, Mycetia, Ophiorrhiza), infrequently differing in shape between long-styled and short-styled forms (Antirhea), infrequently markedly fleshy to leathery (Caelospermum, Damnacanthus, Fosbergia, Rothmannia, Timonius, Urophyllum), inside glabrous to variously pubescent with pubescence frequently confined to throat, outside infrequently ridged to winged (Cinchona, Ophiorrhiza), rarely fenestrate in tube (i.e., with longitudinal slits; Damnacanthus, Paederia), lobes (3 or)4 or 5(–11), shorter than or occasionally longer than tube,