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2. Rubiaceae

茜草科 qian cao ke

Authors: Tao Chen, Xianrui Luo, Hua Zhu, Charlotte M. Taylor, Friedrich Ehrendorfer, Henrik Lantz, Michele Funston & Christian Puff

Galium verum

Credit: Harvard University Herbaria

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 imbricate, 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 occasionally 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 (Hymenodictyon, 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 colleters (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 pseudoaxillary (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 pedicels 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 involucral, 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 protogynous], 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,

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 occasionally 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, acute or less often obtuse to rounded at apex, generally spreading to somewhat reflexed at anthesis, infrequently with margins crisped to irregular, densely ciliate, and/or appendaged (Cinchona, Luculia, Rondeletia, Saprosma, Serissa), in bud imbricate (and usually quincuncial), valvate, valvate-induplicate, valvate-reduplicate, or convolute (i.e., contorted) to left or rarely to right (Coptosapelta, Rothmannia), usually with aestivation consistent within a genus, occasionally with wings, ridges, and/or rounded to hornlike thickenings or protuberances on back or at apex (Lerchea, Ophiorrhiza), rarely cucullate (Lerchea). Stamens adnate to corolla, free or rarely fused to stigma (Acranthera), alternate to corolla lobes and isomerous (i.e., equal in number to lobes) or rarely more numerous (Gardenia) [to rarely fewer], inserted variously in corolla throat, tube, or infrequently at base and sometimes appearing free (Galium), included to exserted, with point of insertion and position of anthers usually differing between long-styled and short-styled forms of distylous flowers, with staminodes of pistillate flowers generally similar to stamens but smaller; filaments well developed to reduced or obsolete, free or rarely coherent (Argostemma) or fused (Acranthera, Argostemma), variously glabrous to pubescent, occasionally markedly flattened (Hymenodictyon, Kelloggia); anthers free or rarely coherent or fused (Argostemma), 4-thecal or rarely 2-thecal (Hymenodictyon), 2-celled, in outline generally narrowly oblong, linear (i.e., narrowly fusiform), narrowly lanceolate, or narrowly elliptic, at base occasionally bifid (i.e., sagittate; Caelospermum, Cephalanthus, Duperrea, Hamelia, Neohymenopogon, Scyphiphora), dorsifixed (i.e., medifixed to dorsifixed near base) to occasionally basifixed, infrequently pubescent (Hyptianthera, Lerchea), with dehiscence introrse by longitudinal slits or rarely by apical pores (Argostemma), with connective infrequently prolonged into an apical and/or sometimes basal appendage (Acranthera, Argostemma, Hyptianthera, Morinda, Rubovietnamia, Wendlandia), appendages rarely fused into a cone (Acranthera); pollen variously 3- or 4(or 5)-colpate and generally subglobose or occasionally 3- or 4-porate, 5-25-colpate and disk-shaped or ellipsoidal, in tetrahedral tetrads and 3- or 4-porate, cylindrical with 2 pores, or inaperturate. Ovary inferior [or rarely secondarily superior], sometimes fused between flowers (Mitchella, Morinda, Mouretia), (1 or)2(-10)-celled (i.e., locular), with ovules 1 or 2 to numerous in each cell (i.e., locule) on basal, axile (i.e., inserted on septum), apical, or infrequently parietal (Ceriscoides, Gardenia) placentas, in staminate flowers usually with ovary reduced and pistillode composed of structures similar to but smaller than style and stigma, in distylous flowers usually with ovary similarly developed but style and stigma differing in size and position in flower and sometimes stigmas also differing in shape between long-styled and short-styled forms; style 1, terminal on ovary, developed or rarely reduced (Galium, Microphysa), variously glabrous to pubescent, surrounded at base by well-developed fleshy disk [or this rarely reduced], this disk variously annular, conical, 2-parted, or shallowly lobed, glabrous or rarely pubescent (Clarkella, Mouretia, Timonius); stigmas free or rarely fused to anther connectives (Acranthera), 1- or 2(-10)-lobed (i.e., these lobes often equivalent to "stigmas 1 or 2 to 10" of some authors), with whole stigma or lobes variously capitate, linear, spatulate, clavate, lobed, or infrequently mitriform (i.e., shaped like a drinking glass) to cylindrical with recessed attachment (Canthium, Mitragyna, Pyrostria), glabrous or infrequently pubescent (Clarkella, Hyptianthera), variously included to exserted, with receptive surfaces introrse, apical, or internal. Infructescences generally similar to inflorescences but occasionally with part or all changing color, orientation, shape, and/or other characteristics as the fruit mature. Fruit simple or rarely multiple (i.e., a syncarp; Mitchella, Morinda, Nauclea), variously capsular with dehiscence loculicidal, septicidal, circumscissile (i.e., around equator or middle; Mitracarpus), or through an apical beak (i.e., prolonged disk portion, sometimes to give appearance of partially superior ovary; Hedyotis, Neanotis, Neohymenopogon, Ophiorrhiza, Pentas, Spiradiclis) or operculum (i.e., circular lid; Argostemma, Leptodermis, Leptomischus, Mouretia, Pseudopyxis); or fleshy, small to quite large, and baccate (i.e., with numerous seeds enclosed by fleshy to juicy pulp or endocarp and usually indehiscent); or drupaceous (i.e., with 1 to several seeds enclosed in pyrenes), fleshy to occasionally dry, and indehiscent or infrequently dehiscent releasing pyrenes (Paederia, Serissa); or schizocarpous (i.e., dry and separating into segments) with mericarps (i.e., segments, cocci, nutlets) indehiscent (e.g., Asperula, Cephalanthus, Richardia); calyx limb persistent or deciduous usually leaving a circular scar, sometimes with persistent carpophore or septum (Adina); pyrenes (i.e., seeds enclosed in and dispersing with endocarp layer) when present and ovules all developed 1-10-locular and 1-10-seeded (i.e., fruit containing several pyrenes and each pyrene with 1 seed in 1 locule, or pyrenes solitary in each fruit and comprising entire ovary), ellipsoid to subglobose, plano-convex (i.e., hemispherical), concavo-convex, lenticular, or angled (i.e., narrow with 2 large inner faces and a small outer face), with outer wall hard to cartilaginous (Caelospermum) or infrequently papery (Coffea, Pavetta), smooth to ridged or sulcate on dorsal (i.e., abaxial) surface, without (Psychotria) or usually with evident preformed germination slits, pores, and/or opercula generally on ventral (i.e., adaxial) surface, rarely winged (Paederia) or pubescent (Caelospermum, Paederia); seeds 1 to numerous, small (0.1-1.9 mm), medium-sized (2-5 mm) to large (5.1-20 mm), variously ellipsoid, lenticular, flattened, oblanceoloid, angled, or plano-convex, smooth to variously winged, foveolate, tuberculate, papillose, and/or striate; endosperm (i.e., albumen) fleshy, oily, corneous (i.e., horny) and entire or infrequently ruminate (Psychotria), or rarely absent (Antirhea); embryo variously shaped. x = 6-17, most commonly 11, less frequently 9 or 12.

About 660 genera and 11,150 species: cosmopolitan family, with most genera and species in humid tropical regions; 97 genera (three endemic, ten introduced) and 701 species (352 endemic, 23 introduced, six of unconfirmed occurrence) in China.

This is one of the largest families of flowering plants and is represented nearly worldwide though it is most abundant in the tropical regions of both hemispheres. Important economic members of the family are coffee (Coffea), quinine (Cinchona), madder (Rubia), ipecac (Carapichea Aublet), and various horticultural plants, notably Gardenia, Ixora, Mussaenda, and Pentas. Several genera include pantropical weeds, notably Mitracarpus, Richardia, and Spermacoce, of neotropical origin, and Oldenlandia (Hedyotis) of African origin. A number of species are used for various medicinal purposes.

Genera, tribes, and subfamilies of Rubiaceae characteristically either have or lack raphides; this character is generally considered informative taxonomically in this family. Raphides are tiny, needle-shaped crystals that are produced in groups inside some cells of leaf, stem, flower, and fruit tissues. They apparently function to protect the plant from herbivores. Raphides are often visible with a hand lens on dried specimens, particularly in soft tissues such as petals, the calyx hypanthium, and the young leaves at the apex of the stem, but may be difficult to see especially in succulent plants.

Leaf domatia are common in Rubiaceae and are sometimes taxonomically informative at least for separation of species. These domatia provide protection for tiny mites that live on the leaves and that help protect the plant from herbivores and fungi by eating insect eggs, small insects, and spores. These domatia occasionally take the form of foveolae or crypts, but more often are formed by small groups or tufts of pubescence, which is sometimes different in form from pubescence found on other parts of the plant.

At least half of the species of Rubiaceae have distylous flowers: an individual species has two flower forms, the long-styled form (or pin) with the stigmas borne above the anthers and the short-styled form (or thrum) with the anthers borne above the stigmas. An individual plant bears only one flower form, and the flowers are incompatible with other flowers of similar form whether borne on the same or another plant. Heterostyly is unusual in the flowering plants and is found in several other plant families that mostly have three floral forms instead of two with the third form (the homostylous) with the stigmas and anthers borne at the same level. Tristyly has only been documented very rarely for Rubiaceae, although it was noted by FRPS for Chassalia curviflora in China.

A number of species of many genera of Rubiaceae have calyx lobes that are unequal in size on an individual flower, and in some species of several genera one or more of the calyx lobes are markedly different, much enlarged and also often petaloid. These structures are found on flowers, where they are typically brightly colored and apparently function to attract pollinators, and/or on the fruit, where they are dry and brown and apparently function in dispersal of the fruit or seeds. These structures have variously been called "petaloid calyx lobes," a morphologically descriptive term; "semaphylls," a term that refers to the assumed function in pollination; and "calycophylls," the term used here that refers to the leaflike shape of the calyx lobe. These structures are sometimes mistaken for bracts; a few species of Rubiaceae do also have petaloid bracts (e.g., Dunnia), but those structures are inserted on the inflorescence axes or immediately below the ovary of the flower.

The woody Rubiaceae are better represented (i.e., more diverse) in moist temperate regions of China than in comparable habitats in North America, as detailed by Latham and Ricklefs (in Ricklefs & Schluter, Spec. Diversity Ecol. Communities, 294-314. 1993). They concluded that this pattern, which is also found in a number of other dicotyledonous families, is due largely to historical factors. The principal factors they identified are the extinction of woody Rubiaceae in these other regions, while the plants were able to persist in E Asia (e.g., Emmenopterys), and the long-term direct connection of the moist temperate and moist tropical zones in E Asia vs. the separation of these zones by deserts and large water bodies in other regions. They agreed with Wolfe (in Nelson & Rosen, Vicariance Biogeogr. 413-427. 1981) that the connections of plant ranges between E North America and E Asia were broken by the end of the Eocene, and thus that species of the several genera that are found in both regions (e.g., Cephalanthus) have had separate evolutionary histories for some time. Several SE Asian Rubiaceae genera, all found in China, appear to represent the most basal living lineages of the family, notably Acranthera, Coptosapelta, and Luculia (Bremer et al., Syst. Biol. 48: 413-435. 1999; Bremer & Manen, Pl. Syst. Evol. 225: 43-72. 2000; Rydin et al., Pl. Syst. Evol. 278: 101-123. 2009).

A number of species of Rubiaceae are cultivated as ornamentals; most of these are mentioned and some are keyed and described here.

Additional information on many of the cultivated Rubiaceae was presented by Puff and Chamchumroon (Thai Forest Bull., Bot. 31: 75-94. 2003).

Complete Rubiaceae floras for E Asia are very few. The Chinese Rubiaceae were treated comprehensively in 1999 by FRPS; the treatment here is based primarily on that work. The Fl. URSS was published in English in 2000, but this is an unedited, unmodified translation of the 1958 work. The Fl. Japan has been published in several editions but includes relatively few genera and is rather idiosyncratic. The genus overview within the Rubiaceae of Thailand: A Pictorial Guide to Indigenous and Cultivated Genera by Puff et al. (43-240. 2005) is a useful recent reference.

Some morphological features that are characteristic of the Rubiaceae have been described, measured, and named very differently by different authors, and the differing terminology has generated some confusion. In particular, the terminology and measurements used for the calyx and gynoecium have differed widely. The ovary of Rubiaceae flowers is typically inferior, and the calyx is gamosepalous with its free portion, or limb, arising from the top of the ovary portion; this limb varies from reduced (i.e., not developed) to developed, and truncate to deeply lobed. Most Rubiaceae authors have distinguished the inferior ovary portion of the flower from the free calyx limb; the ovary portion has sometimes been called simply an ovary, but sometimes considered to include the fused calyx and corolla tissue covering it and called a hypanthium. However, several authors, including in FRPS, have treated as one structure the inferior ovary together with the unlobed lower portion of the calyx limb, when this is present, as the "calyx tube" and the lobed portion of the calyx as a separate structure, the calyx lobes. The "calyx tube" is often clearly delimited and measurable, but the inclusion of the ovary in this means that this measurement can sometimes include ovaries that have been fertilized and are developing into fruit, thus are larger than and not comparable to the ovaries at anthesis even though the corolla may be still attached to the flowers. Here the ovary or hypanthium portion of the flower is measured separately from the limb; the term "hypanthium" is used to link these measurements to those of FRPS. The limb is measured separately here because the length of the tubular portion of this often includes taxonomically useful information, and while the "calyx tube" measurement is lost when the fruit are developed, the length of the tubular portion of the free calyx limb is usually still evident on the fruit thus facilitating identification. Another portion of the gynoecium that has been regarded differently by different authors is the distinction between the stigma and style: the structures regarded as 2-lobed stigmas by some authors are regarded as a style with 2 separate stigmas by as many other authors. Morphological study of the androecium and gynoecium here during preparation of a flora treatment has not been detailed and no focused effort has been made here to systematize these morphological interpretations.

Inflorescence terminology is complicated in most flowering plant families. The terms "bract" and "bracteole" have been carefully and regularly applied by some Rubiaceae authors, with the term "bracteole" used only for the bracts that immediately subtend the flowers or in some cases that are found anywhere along the pedicels, and the term "bract" used for the remaining structures. However, other authors have not distinguished these structures, or have not distinguished them regularly or consistently. The term "floral bract" here refers to the structures found on the pedicels or that immediately subtend sessile flowers. Bracts and leaves are generally considered homologous structures, and in Rubiaceae these often vary continuously along the inflorescences. In particular, Rubiaceae commonly have somewhat reduced leaves subtending the inflorescences, and bracts that are somewhat enlarged and occasionally leaflike. These structures have been variously called "reduced leaves" or "foliose bracts" or "foliaceous bracts" (here called "leaflike bracts"). In particular, many species with branched inflorescences sometimes have rather large, leaflike bracts subtending the basalmost pair of secondary axes; these have been various interpreted as enlarged bracts at the top of the peduncle, or vegetative leaves that are borne at the base of a sessile, tripartite inflorescence.

Significant measurements found on plants from outside China are noted in the discussions following the species descriptions and/or are included within brackets in the descriptions in accordance with Flora of China style. Authors in FRPS described the seeds and embryos of some Rubiaceae but not others; their descriptions are reported. In many cases no information is available about these features for Chinese Rubiaceae. Authors in FRPS also described the pollen of a few genera; these reports are not included here because a number of very detailed, extensively documented pollen descriptions are available elsewhere, which show that much variation is found in pollen morphology at the species level in Rubiaceae and as yet Rubiaceae genera are not well characterized in terms of pollen. Thus, the very limited information available only at the genus level here does not seem well integrated into this flora treatment. The counts presented here for leaf veins apply to the secondary veins, as done by the majority of Rubiaceae authors. Many species of Rubiaceae also frequently have rather well-developed though shorter intersecondary veins, which often vary widely in number and degree of development between populations or plants; these apparently were occasionally combined with the secondary veins in the measurements given by FRPS, but such measurements are subject to significant misinterpretation by authors working in other regions and the varied usage in that work adds yet more confusion.

Where data were missing from the descriptions in the first draft for the Flora of China, the terms "unknown," "not known," "not noted," and "not seen" were retained (with "not known" consisted to "unknown"). Where these terms were used in the habitats and elevational ranges they were deleted in accordance with the style of previous Flora of China volumes. Where missing data were denoted with question marks, e.g., "shape??," or with constructions such as "__ × __ ," these were deleted because it was not apparent which of the above categories of missing data applied.

The genus Khasiaclunea Ridsdale (Blumea 24: 347. 1979) is known with certainty from India and Myanmar and provisionally from N Vietnam; this may also be present in China, but no confidently determined specimens have yet been seen. The single species, K. oligocephala (Haviland) Ridsdale (Blumea 24: 347. 1979; Adina oligocephala Haviland, J. Linn. Soc., Bot. 33: 46. 1897), was reported by Ridsdale (loc. cit.) as "probably" found in China; however, this genus is not treated here.

Lo Hsienshui, Ko Wancheung, Chen Weichiu, Hsue Hsianghao & Wu Hen. 1999. Rubiaceae (1). In: Chen Weichiu, ed., Fl. Reipubl. Popularis Sin. 71(1): i-xvii, 1-432; Lo Hsienshui, Ko Wancheung, Chen Weichiu & Ruan Yunzhen. 1999. Rubiaceae: Rubioideae. In: Chen Weichiu, ed., Fl. Reipubl. Popularis Sin. 71(2): i-xvi, 1-377; Wan Wenhao. 2000. Theligonaceae. In: Chen Chia-jui, ed., Fl. Reipubl. Popularis Sin. 52(2): 147-151.


Key 4

1 Leaves with tertiary or quaternary venation finely, regularly lineolate (this visible best on abaxial surface); inflorescences axillary; fruit a fleshy drupe   (2)
+ Leaves with tertiary and higher order venation not visible or irregularly to somewhat regularly areolate; inflorescences axillary, terminal, or in other positions; fruit dry to fleshy, drupaceous, baccate, capsular, schizocarpous, or other form   (3)
       
2 (1) Fruit with 1 pyrene, this with 2-5 cells (and seeds); corollas 5-11 mm, with 4 or 5 lobes.   5 Antirhea
+ Fruit with 50 or more pyrenes, each with 1 cell; corollas 10-15 mm, with 6 lobes.   92 Timonius
       
3 (1) Plants mangroves, growing along seashores in areas regularly inundated by saltwater, with succulent tissues, usually with stilt roots, usually with young growth covered with mucilage or resin.   83 Scyphiphora
+ Plants of terrestrial or freshwater riverine habitats, with succulent to very thin tissues, without stilt roots, with young growth covered with resin or not   (4)
       
4 (3) Flowers fused together by their ovaries, at least shortly at their bases; fruit multiple (i.e., formed from more than 1 flower)   (5)
+ Flowers free, sometimes tightly packed on a common supporting structure but not actually fused; fruit free (i.e., each formed from only 1 flower)   (6)
       
5 (4) Stipules twisted or imbricate in a hemispherical to conical bud; fruit drupaceous, each with 2 or 4 pyrenes, each pyrene with 1 seed.   54 Morinda
+ Stipules held erect and pressed together in flattened bud; fruit baccate, each fruit with numerous seeds.   59 Nauclea
       
6 (4) Flowers in 1 to several symmetrical, globose heads, heads sessile to pedunculate, terminal or axillary, flowers sessile to pedicellate   (7)
+ Flowers variously arranged in heads, cymes, panicles, or other types of groups, these groups sometimes umbelliform but heads then not symmetrically globose, terminal, axillary, pseudoaxillary, or in other positions   (17)
       
7 (6) Lianas with recurved spines.   94 Uncaria
+ Shrubs or trees, unarmed   (8)
       
8 (7) Some or all leaves truncate to cordate at base   (9)
+ Leaves obtuse to acute at base   (10)
       
9 (8) Inflorescences all axillary; stipules ovate to elliptic or spatulate, unlobed, held erect and pressed together in a flattened bud.   33 Haldina
+ Inflorescences terminal and sometimes also in uppermost leaf axils; stipules ovate in outline and deeply bilobed, imbricate in a conical bud.   85 Sinoadina
       
10 (8) Fruit indehiscent; large trees, frequently cultivated for lumber.   62 Neolamarckia
+ Fruit schizocarpous splitting into 2 mericarps or capsular with septicidal then loculicidal dehiscence; shrubs and trees, perhaps occasionally cultivated   (11)
       
11 (10) Fruit schizocarpous, splitting into 2 indehiscent mericarps; leaves opposite or whorled; shrubs.   13 Cephalanthus
+ Fruit capsular, valves opening to release seeds; leaves opposite; shrubs and trees   (12)
       
12 (11) Stipules held erect and pressed together in a flattened bud   (13)
+ Stipules imbricate or valvate in a conical to hemispherical bud   (14)
       
13 (12) Inflorescence heads sessile or subsessile with peduncles up to 0.5 cm; stigmas cylindrical with style attached in a recessed portion (i.e., upside-down mitriform).   53 Mitragyna
+ Inflorescence heads pedunculate, peduncles 1-8 cm; stigmas globose to ovoid.   63 Neonauclea
       
14 (12) Seeds subglobose to trigonous or ellipsoid, not winged.   49 Metadina
+ Seeds flattened to fusiform or angled, usually winged   (15)
       
15 (14) Calyx lobes ca. 0.5 mm; fruit 5-7 mm.   85 Sinoadina
+ Calyx lobes 1-2 mm; fruit 1.5-5 mm   (16)
       
16 (15) Stipules deeply bilobed for more than 1/2 their length.   2 Adina
+ Stipules triangular and entire to shortly bilobed for 1/4 or less their length.   69 Pertusadina
       
17 (6) Fruit capsular, opening through an apical operculum and later sometimes splitting into 5 valves, with each seed individually enclosed in a fibrous, netlike aril.   44 Leptodermis
+ Fruit indehiscent, baccate, drupaceous, or capsular but then not operculate and with fewer than 5 valves, seeds without aril or with aril continuous, fleshy   (18)
       
18 (17) Inflorescences not borne at stem apex but regularly axillary (i.e., borne in both axils at a node), superaxillary, and/or apparently axillary due to regularly produced axillary or lateral short shoots (i.e., brachyblasts)   (19)
+ Inflorescences terminal, pseudoaxillary (i.e., regularly borne in only 1 axil at a node), or apparently pseudoaxillary or lateral due to anisophylly or an undeveloped internode   (41)
       
19 (18) Stems and abaxial surface of leaf densely sericeous, hirsute, or strigose with remarkably long silky trichomes.   97 Xanthophytum
+ Stems and abaxial surface of leaf variously glabrous to pubescent with trichomes of various types, but not densely and consistently long and silky   (20)
       
20 (19) Inflorescence borne on regular paired axillary short shoots (i.e., brachyblasts), each short shoot with several stipulate nodes; fruit fleshy and drupaceous or baccate, or dry and indehiscent or schizocarpous   (21)
+ Inflorescences borne on peduncles directly from main stems, these with no more than 1 reduced basal node   (31)
       
21 (20) Stipules with 1-8 stiffly spiniform bristles.   84 Serissa
+ Stipules entire or with several to numerous bristles or projections but these not spiny   (22)
       
22 (21) Bracts multifid with segments glandular   (23)
+ Bracts absent or entire and not glandular   (24)
       
23 (22) Corolla lobes valvate in bud; fruit a fleshy drupe.   20 Damnacanthus
+ Corolla lobes convolute in bud; fruit a dry schizocarp.   93 Trailliaedoxa
       
24 (22) Stipules bilobed.   72 Prismatomeris
+ Stipules triangular   (25)
       
25 (24) Flowers with 4 calyx lobes, corolla lobes, and stamens   (26)
+ Flowers with 5 calyx lobes, corolla lobes, and stamens   (28)
       
26 (25) Petioles and inflorescence axes articulate at their bases; stigmas fusiform.   39 Ixora
+ Petioles and inflorescence axes not articulate; stipules lobed   (27)
       
27 (26) Plants without fetid odor; corolla salverform, with lobes convolute in bud.   67 Pavetta
+ Plants with or without fetid odor when bruised; corolla campanulate, tubular, or funnelform, with lobes valvate or valvate-induplicate in bud.   81 Saprosma
       
28 (25) Calyx lobes 5-8 mm.   12 Catunaregam
+ Calyx lobes 0.5-4 mm   (29)
       
29 (28) Ovules and seeds 1-4 in each cell; plants without spines; flowers sessile and solitary.   36 Himalrandia
+ Ovules and seeds more than 4 in each cell; plants with or without spines; flowers sessile to pedicellate, solitary to several in fascicles   (30)
       
30 (29) Plants with bisexual flowers; ovary with ovules and seeds attached to septum (i.e., axile).   8 Benkara
+ Plants dioecious; ovary with ovules and seeds attached to outer wall (i.e., parietal); nodes of stems mostly reduced.   14 Ceriscoides
       
31 (20) Inflorescences racemiform or spiciform, with well-developed primary axes directly bearing numerous pedicels.   37 Hymenodictyon
+ Inflorescences capitate to cymose, without well-developed primary axes or with axes of several orders developed, with higher order axes or all axes bearing few to numerous pedicels   (32)
       
32 (31) Woody twiners or climbers; fruit a woody capsule, subglobose, with winged seeds.   19 Coptosapelta
+ Erect shrubs and trees; fruit fleshy and indehiscent or capsular, papery to cartilaginous, flattened, with unwinged seeds   (33)
       
33 (32) Fruit capsular, papery to cartilaginous, mitriform and laterally markedly flattened (this shape often evident in young developing fruit).   65 Ophiorrhiza
+ Fruit indehiscent and fleshy   (34)
       
34 (33) Corolla lobes imbricate in bud; fruit a drupe with a single, 4-9-celled subglobose pyrene, this 2-3 cm in diam.; plants of coastal thickets.   31 Guettarda
+ Corolla lobes convolute or valvate in bud or imbricate in some species of Lasianthus; fruit baccate with numerous seeds or drupaceous with 1-celled pyrenes 2 to numerous; plants of interior habitats   (35)
       
35 (34) Cultivated plants; corolla lobes convolute in bud; fruit drupaceous with 2 plano-convex pyrenes with papery walls; bracts fused in calyculate pairs.   18 Coffea
+ Native plants; corolla lobes convolute, valvate, or imbricate in bud; fruit baccate or drupaceous with 2 to numerous plano-convex to angled, ellipsoid-oblong, ellipsoid, or flattened, hard-walled pyrenes; bracts free or fused in calyculate pairs   (36)
       
36 (35) Fruit a berry with several to numerous seeds   (37)
+ Fruit a drupe, with 2 to numerous pyrenes, each containing 1 seed   (39)
       
37 (36) Seeds numerous; corolla lobes valvate in bud.   95 Urophyllum
+ Seeds 3-8; corolla lobes convolute in bud   (38)
       
38 (37) Ovules and seeds 1-3 per cell; stigmas papillose; calyx lobes minute or up to 0.5 mm.   23 Diplospora
+ Ovules and seeds 3-6 per cell; stigmas pubescent; calyx lobes 1-2 mm.   38 Hyptianthera
       
39 (36) Flower buds acute to rounded at apex; pyrenes flattened to triangular; plants often with fetid odor when bruised.   43 Lasianthus
+ Flower buds sharply acute to acuminate at apex; pyrenes plano-convex to ellipsoid-oblong or ellipsoid; plants without fetid odor   (40)
       
40 (39) Plants without or sometimes with spines; stipules pilose to sericeous inside (i.e., adaxially).   11 Canthium
+ Plants without spines; stipules glabrous inside.   75 Psydrax
       
41 (18) Stipules with 1-8 sharp spines or spiny bristles.   84 Serissa
+ Stipules entire to lobed or multifid, sometimes aristate or caudate but not spiny   (42)
       
42 (41) Stipules with several well-developed, fibrous, closely set veins, these parallel to palmate.   58 Myrioneuron
+ Stipules smooth, 1-3-costate but not densely veined   (43)
       
43 (42) Calyx limb with developed truncate tube; flowers several to numerous, all pedicellate, borne in umbelliform cymules.   10 Caelospermum
+ Calyx limb reduced to developed and at least shortly lobed or denticulate; flowers variously arranged but not uniformly pedicellate in umbelliform cymules   (44)
       
44 (43) Corolla lobes convolute in bud and fruit a berry with fleshy to leathery outer wall   (45)
+ Corolla lobes valvate, valvate-induplicate, imbricate, or convolute in bud and fruit a drupe, berry, capsule, or schizocarp, if corolla lobes convolute then fruit not a berry   (60)
       
45 (44) Fruit and usually ovary portion of calyx 5-8-ridged or -winged, ridges continuous with midrib of each calyx lobe.   29 Gardenia
+ Fruit and ovary portion of calyx smooth to tuberculate   (46)
       
46 (45) Some or all inflorescences regularly borne in pseudoaxillary, leaf-opposed, and/or lateral position with only 1 inflorescence per node (i.e., not paired, borne on only one side of stem), this arrangement due to anisophylly or a reduced internode producing apparently alternate leaves or a 3-leaved node   (47)
+ Inflorescence borne at stem apices and sometimes in adjacent uppermost leaf axils, sometimes displaced to pseudoaxillary by subsequent overtopping stem growth, produced at stems generally with number and arrangement of leaves similar at all nodes   (51)
       
47 (46) Inflorescences all apparently leaf-opposed or borne at leafless nodes (due to extreme anisophylly), stems with paired or apparently alternate leaves.   3 Aidia
+ Inflorescences mostly borne on apparently 3-leaved nodes, stems with some 3-leaved nodes   (48)
       
48 (47) Flowers 12 or more per inflorescence.   25 Duperrea
+ Flowers 1-10 per inflorescence   (49)
       
49 (48) Some or all leaf apices with short filamentous arista or projection.   80 Rubovietnamia
+ Leaf apices without filamentous projections   (50)
       
50 (49) Plants dioecious, with unisexual flowers; fruit 5-15 mm in diam.; corollas (so far as known) 5-8 mm.   9 Brachytome
+ Plants with bisexual flowers; fruit ca. 35 mm in diam.; corollas 68-75 mm.   78 Rothmannia
       
51 (46) Low shrubs, up to 0.5 m tall, with many internodes reduced and relatively small leaves   (52)
+ Shrubs, low (ca. 0.3 m tall) to tall, or trees   (53)
       
52 (51) Flowers solitary.   36 Himalrandia
+ Flowers in cymes of 6-12.   93 Trailliaedoxa
       
53 (51) Corolla densely sericeous (i.e., with long, silky, appressed trichomes) on outside.   71 Porterandia
+ Corolla glabrous to puberulent, spreading pubescent, or strigillose (i.e., with short appressed trichomes) on outside   (54)
       
54 (53) Inflorescences with 1-7 flowers; fruit 4-11 cm.   27 Fosbergia
+ Inflorescences with 5 to numerous flowers; fruit 2 cm or shorter   (55)
       
55 (54) Petioles and inflorescence axes articulate; ovules and seeds 1 per cell, 2 per ovary or fruit; stigmas unlobed.   39 Ixora
+ Petioles and inflorescence axes not articulate; ovules and seeds 1 to numerous per cell, 2 to numerous per ovary or fruit; stigmas lobed   (56)
       
56 (55) Bark of older stems becoming coppery red and circumferentially wrinkled to peeling.   90 Tarennoidea
+ Bark of older stems smooth, persistent, green to gray or brown, sometimes becoming longitudinally striate or fissured with age   (57)
       
57 (56) Ovules 1 per cell and 2 per ovary; fruit drupaceous with 2 plano-convex pyrenes.   67 Pavetta
+ Ovules 2 or more per cell and 4 or more per ovary; fruit baccate with seeds 2 or more, ellipsoid, obovoid, or compressed globose   (58)
       
58 (57) Seeds 2-6 per fruit, 4-5 mm in diam., compressed globose, fruit yellowish white; plants without spines.   4 Alleizettella
+ Seeds 4 to numerous per fruit, 3-5 mm in diam., compressed globose to ellipsoid or obovoid, fruit purplish black, black, or sometimes white; plants with or without spines   (59)
       
59 (58) Plants without or usually with spines; calyx limb persistent or deciduous on fruit.   8 Benkara
+ Plants without spines; calyx limb deciduous on fruit.   89 Tarenna
       
60 (44) Stigmas, cells of ovary, and cells of fruit 4 or 5   (61)
+ Stigmas 1 or 2 and cells of ovary and fruit 2, or sometimes any of these 3 on some but not all flowers in an inflorescence   (64)
       
61 (60) Corollas tubular with relatively short lobes, these 10% or less of total corolla length; plants cultivated ornamentals.   34 Hamelia
+ Corollas funnelform, tubular-funnelform, or salverform with lobes developed, 15% or more of total corolla length; plants native or cultivated   (62)
       
62 (61) Fruit dry, schizocarpous or capsular, leathery; plants often with fetid odor when bruised.   87 Spermadictyon
+ Fruit fleshy and drupaceous or baccate, or capsular and thinly textured; plants without fetid odor   (63)
       
63 (62) Plants not particularly succulent, sometimes with spines, usually with some internodes not expanded; calyx without stalked glands.   20 Damnacanthus
+ Plants sometimes succulent, without spines, with all internodes expanded; calyx sometimes with stalked glands.   57 Mycetia
       
64 (60) Cultivated ornamental plants; herbs and low shrubs to small trees; corollas red, orange, or yellow   (65)
+ Native plants; low shrubs to large trees; corollas white, pink, purple, yellow, or red   (68)
       
65 (64) Stigmas unlobed; corolla lobes convolute in bud; petioles and inflorescence axes articulate.   39 Ixora
+ Stigmas 2- or 3-lobed; corolla lobes valvate, valvate-induplicate, or imbricate in bud; petioles and inflorescence axes not articulate   (66)
       
66 (65) Plants often with petaloid calycophylls; corolla lobes valvate-induplicate in bud.   56 Mussaenda
+ Plants without petaloid calycophylls; corolla lobes valvate or imbricate in bud   (67)
       
67 (66) Herbs and low shrubs; corolla with white pubescence in throat, lobes obtuse to acute, with margins flat.   68 Pentas
+ Shrubs; corolla with yellow pubescence in throat, lobes broadly obtuse to rounded, with margins flat to crisped.   77 Rondeletia
       
68 (64) Fruit indehiscent, drupaceous, or baccate, fleshy or leathery; corolla lobes convolute, imbricate, or valvate in bud; ovules and seeds 1 to numerous per cell, 2 to numerous per fruit   (69)
+ Fruit dry and capsular (i.e., dehiscent), indehiscent, or schizocarpous (i.e., splitting into sections); corolla lobes valvate, valvate-induplicate, or imbricate in bud; ovules and seeds several to numerous in each cell and fruit, or 1 per cell and 2 per fruit in Trailliaedoxa   (79)
       
69 (68) Stigmas unlobed; petioles and inflorescence axes articulate at base.   39 Ixora
+ Stigmas 2- or 3-lobed; petioles and inflorescence axes not articulate   (70)
       
70 (69) Fruit baccate, with numerous seeds in each cell, these enclosed in soft tissues   (71)
+ Fruit indehiscent or drupaceous, with 1 seed in each cell, this enclosed in a pyrene   (72)
       
71 (70) Corolla lobes valvate-reduplicate; calyx sometimes with calycophylls, without stalked glands; seeds flattened; plants not succulent.   56 Mussaenda
+ Corolla lobes valvate-induplicate; calyx without calycophylls, sometimes with stalked glands; seeds angled; plants often succulent.   57 Mycetia
       
72 (70) Stipules caducous or rather quickly deciduous, often exposing a ring of persistent trichomes that usually dry red-brown (use magnification).   74 Psychotria
+ Stipules persistent to caducous, not enclosing persistent trichomes or these reduced and/or drying white   (73)
       
73 (72) Corolla lobes convolute in bud   (74)
+ Corolla lobes valvate in bud   (75)
       
74 (73) Fruit drupaceous, with pyrenes flattened to concavo-convex.   67 Pavetta
+ Fruit schizocarpous with segments or valves oblanceoloid.   93 Trailliaedoxa
       
75 (73) Corolla tubular-funnelform with base usually gibbous and tube usually curved.   15 Chassalia
+ Corolla tubular-funnelform or salverform, generally straight at base and in tube   (76)
       
76 (75) Stipules persistent and becoming hardened and yellowed with age, often fragmenting when older.   74 Psychotria
+ Stipules caducous, or persistent but then not becoming hardened, yellowed, or fragmented   (77)
       
77 (76) Plants often with fetid odor when bruised; corolla lobes mostly 4, or on a few flowers of some plants 5 or 6.   81 Saprosma
+ Plants without fetid odor; corolla lobes regularly 5, on a few flowers of some plants 4   (78)
       
78 (77) Bracts glandular-multifid.   20 Damnacanthus
+ Bracts entire, not glandular.   72 Prismatomeris
       
79 (68) Inflorescences racemiform or spiciform, with 1 or a few well-developed main axes bearing pedicels.   37 Hymenodictyon
+ Inflorescences subcapitate to cymose, with axes of various orders developed to different degrees and flowers usually borne from axes of various orders   (80)
       
80 (79) Fruit mitriform and laterally markedly flattened (this shape often evident in young developing fruit).   65 Ophiorrhiza
+ Fruit ellipsoid to subglobose   (81)
       
81 (80) Fruit capsular, septicidal, 8-50 mm with at least some fruit 10 mm or longer; seeds flattened and often winged; corollas 5-70 mm   (82)
+ Fruit indehiscent, schizocarpous, or capsular, septicidal or loculicidal, 1-8 mm; seeds flattened and sometimes winged to angled and unwinged; corollas 2-20 mm   (85)
       
82 (81) Stipules persistent; shrubs, often epiphytic.   61 Neohymenopogon
+ Stipules caducous; terrestrial shrubs or trees   (83)
       
83 (82) Corolla lobes valvate in bud, acute; cultivated trees, without petaloid calycophylls.   16 Cinchona
+ Corolla lobes imbricate in bud, obtuse to rounded; native trees perhaps sometimes cultivated, sometimes with petaloid calycophylls   (84)
       
84 (83) Calyx lobes 2-2.5 mm; corolla lobes ca. 7 mm; plants sometimes with calycophylls.   26 Emmenopterys
+ Calyx lobes 8-20 mm; corolla lobes 11-18 mm; plants without calycophylls.   48 Luculia
       
85 (81) Corolla lobes imbricate in bud; fruit capsular, loculicidal, subglobose, woody.   96 Wendlandia
+ Corolla lobes valvate, valvate-induplicate, or convolute in bud; fruit indehiscent, schizocarpous, or capsular and septicidal or loculicidal, subglobose to ellipsoid, papery to woody, if loculicidal then corolla lobes valvate   (86)
       
86 (85) Corolla lobes convolute in bud; low shrubs; ovules and seeds 1 per cell.   93 Trailliaedoxa
+ Corolla lobes valvate or valvate-induplicate in bud; low to large shrubs; ovules and seeds several to numerous in each cell   (87)
       
87 (86) Corolla lobes valvate-induplicate in bud; plants often with petaloid calycophylls.   82 Schizomussaenda
+ Corolla lobes [simple-]valvate in bud; plants with or without petaloid calycophylls   (88)
       
88 (87) Inflorescences usually with 1 or several petaloid bracts and/or calycophylls; corolla yellow; capsules septicidal; stipules interpetiolar, persistent.   24 Dunnia
+ Inflorescences without petaloid bracts or calycophylls; corolla white, pink, blue, purple, or yellow; capsules septicidal or loculicidal or fruit indehiscent; stipules interpetiolar or fused to petiole bases, persistent or caducous   (89)
       
89 (88) Fruit dehiscent through apical part or splitting into 2 indehiscent valves.   35 Hedyotis
+ Fruit deeply dehiscent into 2 or 4 separate valves.   88 Spiradiclis

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