The World Flora Online

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<

Trees, shrubs or rarely herbs Leaves opposite or rarely verticillate, entire; stipules inter- or intra-petiolar, often connate, rarely leafy and not distinguishable from the leaves Seeds rarely winged, mostly with endosperm, the latter rarely ruminate; embryo straight or curved Ovary inferior or rarely superior, 2- or more-celled, with axile, apical or basal placentas, or rarely 1-celled with parietal placentas; style often slender; ovules 1 to many Fruit a capsule, berry or drupe Corolla epigynous, more or less tubular, rarely campanulate; lobes 4-12, contorted, imbricate or valvate Stamens epipetalous, as many as and alternate with the corolla-lobes; anthers mostly separate, 2-celled, opening lengthwise or rarely by terminal pores, rarely transversely septate Flowers usually hermaphrodite, actinomorphic or very rarely slightly zygomorphic, solitary to capitate Calyx adnate to the ovary Leaves opposite or rarely verticillate, entire; stipules inter- or intra-petiolar, often connate, rarely leafy and not distinguishable from the leaves Seeds rarely winged, mostly with endosperm, the latter rarely ruminate; embryo straight or curved Ovary inferior or rarely superior, 2- or more-celled, with axile, apical or basal placentas, or rarely 1-celled with parietal placentas; style often slender; ovules 1 to many Fruit a capsule, berry or drupe Corolla epigynous, more or less tubular, rarely campanulate; lobes 4-12, contorted, imbricate or valvate Stamens epipetalous, as many as and alternate with the corolla-lobes; anthers mostly separate, 2-celled, opening lengthwise or rarely by terminal pores, rarely transversely septate Flowers usually hermaphrodite, actinomorphic or very rarely slightly zygomorphic, solitary to capitate Calyx adnate to the ovary

Fls perfect, mostly 4(–8)-merous, epigynous, sympetalous, regular; cal mostly small, with open aestivation, or obsolete; stamens on the cor-tube, as many as and alternate with the lobes; ovary inferior, mostly 2(–5)-carpellate and plurilocular, with axile (or axile-basal or axile-apical) placentation; style slender, terminal, with lobed or capitate stigma, or (as in Galium) the styles distinct; ovules and seeds 1–many per locule; seeds with a dicotyledonous, usually straight embryo embedded in the oily endosperm; herbs or more often woody plants with simple, mostly entire, commonly decussately opposite lvs and interpetiolar (usually connate) stipules, or with whorled lvs and no stipules, the stipules typically bearing colleters within, or seldom the stipules reduced to mere interpetiolar lines; infls basically cymose. 450/6500, cosmop. but mostly trop. and subtrop.

Flowers rarely solitary, mostly in various terminal or axillary inflorescences, all basically cymose but variously aggregated into panicles, etc., occasionally in globose heads to an extent that the ovaries are adnate; bracts vestigial to well developed, even conspicuous; flowers usually hermaphrodite, rarely unisexual, regular or nearly so (except in >i>Posoqueria (America)) or corolla-tube rarely curved, homostylous or quite often heterostylous with 2 or rarely 3 forms (long-styled (dolichostylous), short-styled (brachystylous) or equal-styled (isostylous)) Calyx gamosepalous, the tube mostly adnate to the ovary, (3–)4–5(–8)-toothed or -lobed, sometimes only minutely so, with open, valvate, imbricate or contorted aestivation, 1 or several lobes sometimes slightly to very considerably enlarged to form a leafy often coloured lamina Small to large trees, shrubs or less often annual or perennial herbs or woody or herbaceous climbers, sometimes spiny; tissues in many tribes containing abundant rhaphides Leaves opposite or verticillate, decussate, almost always entire, very rarely (not in East Africa) palmatifid, toothed or finely denticulate, always obviously stipulate (save in some Rubieae, where the stipules may be considered foliar or almost absent according to interpretation), the stipules interpetiolar or intrapetiolar, entire or, particularly in herbaceous genera, variously divided into lobes or fimbrieae, often tipped or separated by mucilaginous hairs known as colleters and often with colleters inside the base; the leaves in certain tribes sometimes contain small bacterial nodules Corolla small to large and showy, gamopetalous, rotate to salver-shaped or funnel-shaped, the tube often very long, (3–)4–5(–11)-lobed, the lobes mostly contorted or valvate, sometimes valvate-induplicate, rarely imbricate or quincuncial Stamens usually as many as the corolla-lobes and alternate with them, epipetalous; anthers basi- or dorsifixed, introrse, the thecae rarely multilocellate transversely Pollen various, mostly simple, isopolar and 3-colporate, but sometimes porate, the number of colpi or pores varying from 2 to 25, globose, ovoid or discoid, sometimes (in some >i>Gardenieae) in tetrads or rarely in polyads Disc often present, 2-lobed or tubular Ovary inferior, rarely half-inferior or (in >i>Gaertnera) superior, syncarpous of 2–5 or more carpels, but predominantly of 2 and therefore predominantly 2-locular, but 3–5 or even 12 or more (e.g. in >i>Urophylleae due to supplementary incomplete partitions); placentation axile or (in some >i>Gardenieae) parietal; ovules 1–many per locule, often embedded in fleshy placentas, erect, basal or horizontal, anatropous; style simple, usually long and narrow, the “stigma” either cylindrical, clavate, or otherwise modified to form a “receptaculum pollinis” or divided into 2–many linear, spathulate or clavate lobes, the actual stigmatic surface sometimes confined to certain areas, e.g. the inner faces of the lobes Fruit small to quite large (0.2–20 cm.), a capsule, berry or drupe or indehiscent or woody, occasionally (e.g. in >i>Nauclea and >i>Morinda) united to form syncarps, (1–)2–many-seeded, if capsules then loculicidal or septicidal or opening by a beak Seeds small to rather large, sometimes winged; testa cells in some tribes with very distinct pits; albumen present (save in >i>Guettardeae); embryo straight or rarely curved, the radicle mostly longer than the cotyledons Calyx gamosepalous, the tube mostly adnate to the ovary, (3–)4–5(–8)-toothed or -lobed, sometimes only minutely so, with open, valvate, imbricate or contorted aestivation, 1 or several lobes sometimes slightly to very considerably enlarged to form a leafy often coloured lamina Small to large trees, shrubs or less often annual or perennial herbs or woody or herbaceous climbers, sometimes spiny; tissues in many tribes containing abundant rhaphides Leaves opposite or verticillate, decussate, almost always entire, very rarely (not in East Africa) palmatifid, toothed or finely denticulate, always obviously stipulate (save in some Rubieae, where the stipules may be considered foliar or almost absent according to interpretation), the stipules interpetiolar or intrapetiolar, entire or, particularly in herbaceous genera, variously divided into lobes or fimbrieae, often tipped or separated by mucilaginous hairs known as colleters and often with colleters inside the base; the leaves in certain tribes sometimes contain small bacterial nodules Corolla small to large and showy, gamopetalous, rotate to salver-shaped or funnel-shaped, the tube often very long, (3–)4–5(–11)-lobed, the lobes mostly contorted or valvate, sometimes valvate-induplicate, rarely imbricate or quincuncial Stamens usually as many as the corolla-lobes and alternate with them, epipetalous; anthers basi- or dorsifixed, introrse, the thecae rarely multilocellate transversely Pollen various, mostly simple, isopolar and 3-colporate, but sometimes porate, the number of colpi or pores varying from 2 to 25, globose, ovoid or discoid, sometimes (in some >i>Gardenieae) in tetrads or rarely in polyads Disc often present, 2-lobed or tubular Ovary inferior, rarely half-inferior or (in >i>Gaertnera) superior, syncarpous of 2–5 or more carpels, but predominantly of 2 and therefore predominantly 2-locular, but 3–5 or even 12 or more (e.g. in >i>Urophylleae due to supplementary incomplete partitions); placentation axile or (in some >i>Gardenieae) parietal; ovules 1–many per locule, often embedded in fleshy placentas, erect, basal or horizontal, anatropous; style simple, usually long and narrow, the “stigma” either cylindrical, clavate, or otherwise modified to form a “receptaculum pollinis” or divided into 2–many linear, spathulate or clavate lobes, the actual stigmatic surface sometimes confined to certain areas, e.g. the inner faces of the lobes Fruit small to quite large (0.2–20 cm.), a capsule, berry or drupe or indehiscent or woody, occasionally (e.g. in >i>Nauclea and >i>Morinda) united to form syncarps, (1–)2–many-seeded, if capsules then loculicidal or septicidal or opening by a beak Seeds small to rather large, sometimes winged; testa cells in some tribes with very distinct pits; albumen present (save in >i>Guettardeae); embryo straight or rarely curved, the radicle mostly longer than the cotyledons

Seeds small to rather large, sometimes winged; testa cells in some tribes with very distinct pits; endosperm present (save in Guettardeae), entire or ruminate; embryo straight or rarely curved; radicle erect or horizontal, mostly longer than the cotyledons, orientation pendulous Fruit small to very large, 2–many-seeded, dehiscent (capsules either loculicidal, septicidal, opening by a beak or infrequently circumscissile, or splitting into mericarps), or indehiscent (berries, including forms with woody walls, or drupes), occasionally (eg in Nauclea and Morinda) united to form syncarps; if drupes containing one-seeded pyrenes thin- or thick-walled, opening by pre-formed germination slits or not, or less often a multi-seeded stone Ovary inferior, rarely half-inferior or superior (in Gaertnera), syncarpous of 2–10(12 or more) carpels, but predominantly of 2, and therefore predominantly 2-locular, but 3–10(12) (in Urophylleae can appear higher than actual carpel number due to supplementary incomplete partitions), or infrequently uni-locular (or partly so); ovules l–numerous per locule; placentation axile (attached to septum at apex, middle or base, usually on a placenta, often embedded in fleshy placentas) or infrequently (in some Gardenieae) parietal; ovules anatropous; style simple, usually long (but very short in Rubieae) and narrow, at the top either cylindrical, clavate, or otherwise modified to form a pollen presenter (the actual stigmatic surface sometimes confined to specific areas, eg the inner faces of the lobes**), or divided into 2–many (according to number of carpels) linear, spathulate or clavate stigmatic lobes Disk usually functioning as a nectary, positioned above the ovary inside the base of the calyx limb, annular to cushion-shaped or tubular, sometimes 2-lobed, occasionally fused to base of calyx limb-tube (Naucleeae) or absent (in wind-pollinated flowers) Pollen various, mostly simple, isopolar and 3- colporate, but sometimes porate, the number of colpi or pores varying from 2 to 25, globose, ovoid or discoid, sometimes (in some Gardenieae) in tetrads or rarely in polyads Stamens usually as many as the corolla lobes and alternate with them, epipetalous or not (some New World tribes only); anthers basi- or more often dorsifixed, introrse rarely porate (not in the Flora Zambesiaca area), the thecae rarely transversely multilocellate Corolla predominantly white but sometimes coloured, small to large and showy, gamopetalous, mostly salver- shaped to funnel-shaped or less often campanulate, occasionally tube very short, the tube rarely curved, (3)4–5(11)-lobed, the lobes contorted (mostly to the left, infrequently to the right) or valvate (sometimes induplicate or reduplicate), with or without a contact zone (i.e. thickened margin), less often imbricate (tab. 73/B–D) Calyx gamosepalous, the tube (i.e. calyx tube**) mostly adnate to the ovary, the limb with tubular part (i.e. limb-tube) present or absent, truncate or bearing minute teeth to well developed lobes (tab. 73/A1–A4), valvate (often separated by sinuses), or less often imbricate or contorted in aestivation, in a few genera 1–several lobes sometimes slightly to considerably enlarged to form a leaf-like often coloured blade (calycophyll) (tab. 73/A5) Stipules interpetiolar or occasionally intrapetiolar (mostly New World taxa); lobes shortly fused above the petioles and sometimes forming a sheath, or lobes free, entire, bifid or rarely laciniate, or particularly in herbaceous genera, fimbriate, often tipped with or separated by colleters (mucilaginous glands) (tab. 74), often with colleters inside the limb, especially towards the base Inflorescences terminal or axillary, basically thyrsoid, variously aggregated into panicles etc., occasionally in globose heads sometimes to an extent that the ovaries are adnate, or sometimes flowers solitary; bracts vestigial to well developed, even conspicuous Flowers regular or slightly to distinctly (not in Africa) zygomorphic, hermaphrodite or less often unisexual, homostylous, in which case very often protandrous, with or without pollen presentation (tab. 73/F,G), or rarely protogynous (not in Africa), or heterostylous with 2 or less often 3 forms (long-styled (dolichostylous), short-styled (brachystylous) or equal-styled (isostylous) (tab. 73/E) Small to large trees, shrubs (including suffrutices from woody rootstocks) or less often annual or perennial herbs or woody or herbaceous climbers, sometimes spiny; raphides (tab. 1/7) present or absent; crystal sand absent or present (these two mutually exclusive) Leaves opposite or occasionally verticillate, decussate, almost always entire, very rarely sparsely toothed or lobed (not in the Flora Zambesiaca area), always obviously stipulate (save in some Rubieae, where the stipules may be considered leaf-like or almost absent according to interpretation); domatia (mite associated pits or tufts of hairs) very often present axils of lateral nerves on the leaf lower surface; bacterial nodules present in a few genera Flowers regular or slightly to distinctly (not in Africa) zygomorphic, hermaphrodite or less often unisexual, homostylous, in which case very often protandrous, with or without pollen presentation (tab. 73/F,G), or rarely protogynous (not in Africa), or heterostylous with 2 or less often 3 forms (long-styled (dolichostylous), short-styled (brachystylous) or equal-styled (isostylous) (tab. 73/E) Corolla predominantly white but sometimes coloured, small to large and showy, gamopetalous, mostly salver- shaped to funnel-shaped or less often campanulate, occasionally tube very short, the tube rarely curved, (3)4–5(11)-lobed, the lobes contorted (mostly to the left, infrequently to the right) or valvate (sometimes induplicate or reduplicate), with or without a contact zone (i.e. thickened margin), less often imbricate (tab. 73/B–D) Calyx gamosepalous, the tube (i.e. calyx tube**) mostly adnate to the ovary, the limb with tubular part (i.e. limb-tube) present or absent, truncate or bearing minute teeth to well developed lobes (tab. 73/A1–A4), valvate (often separated by sinuses), or less often imbricate or contorted in aestivation, in a few genera 1–several lobes sometimes slightly to considerably enlarged to form a leaf-like often coloured blade (calycophyll) (tab. 73/A5) Pollen various, mostly simple, isopolar and 3- colporate, but sometimes porate, the number of colpi or pores varying from 2 to 25, globose, ovoid or discoid, sometimes (in some Gardenieae) in tetrads or rarely in polyads Stamens usually as many as the corolla lobes and alternate with them, epipetalous or not (some New World tribes only); anthers basi- or more often dorsifixed, introrse rarely porate (not in the Flora Zambesiaca area), the thecae rarely transversely multilocellate Ovary inferior, rarely half-inferior or superior (in Gaertnera), syncarpous of 2–10(12 or more) carpels, but predominantly of 2, and therefore predominantly 2-locular, but 3–10(12) (in Urophylleae can appear higher than actual carpel number due to supplementary incomplete partitions), or infrequently uni-locular (or partly so); ovules l–numerous per locule; placentation axile (attached to septum at apex, middle or base, usually on a placenta, often embedded in fleshy placentas) or infrequently (in some Gardenieae) parietal; ovules anatropous; style simple, usually long (but very short in Rubieae) and narrow, at the top either cylindrical, clavate, or otherwise modified to form a pollen presenter (the actual stigmatic surface sometimes confined to specific areas, eg the inner faces of the lobes**), or divided into 2–many (according to number of carpels) linear, spathulate or clavate stigmatic lobes Disk usually functioning as a nectary, positioned above the ovary inside the base of the calyx limb, annular to cushion-shaped or tubular, sometimes 2-lobed, occasionally fused to base of calyx limb-tube (Naucleeae) or absent (in wind-pollinated flowers) Seeds small to rather large, sometimes winged; testa cells in some tribes with very distinct pits; endosperm present (save in Guettardeae), entire or ruminate; embryo straight or rarely curved; radicle erect or horizontal, mostly longer than the cotyledons, orientation pendulous Fruit small to very large, 2–many-seeded, dehiscent (capsules either loculicidal, septicidal, opening by a beak or infrequently circumscissile, or splitting into mericarps), or indehiscent (berries, including forms with woody walls, or drupes), occasionally (eg in Nauclea and Morinda) united to form syncarps; if drupes containing one-seeded pyrenes thin- or thick-walled, opening by pre-formed germination slits or not, or less often a multi-seeded stone Stipules interpetiolar or occasionally intrapetiolar (mostly New World taxa); lobes shortly fused above the petioles and sometimes forming a sheath, or lobes free, entire, bifid or rarely laciniate, or particularly in herbaceous genera, fimbriate, often tipped with or separated by colleters (mucilaginous glands) (tab. 74), often with colleters inside the limb, especially towards the base Inflorescences terminal or axillary, basically thyrsoid, variously aggregated into panicles etc., occasionally in globose heads sometimes to an extent that the ovaries are adnate, or sometimes flowers solitary; bracts vestigial to well developed, even conspicuous Small to large trees, shrubs or less often annual or perennial herbs or woody or herbaceous climbers, sometimes spiny; tissues in many tribes containing abundant rhaphides Leaves opposite or verticillate, decussate, almost always entire, very rarely (not in Africa) palmatifid, toothed or finely denticulate, always obviously stipulate (save in some Rubieae, where the stipules may be considered foliar or almost absent according to interpretation), the stipules interpetiolar or intrapetiolar, entire or, particularly in herbaceous genera, variously divided into lobes or fimbriae, often tipped or separated by mucilaginous hairs known as colleters and often with colleters inside the base; the leaves in certain tribes sometimes contain small bacterial nodules Corolla small to large and showy, gamopetalous, rotate to salver-shaped or funnel-shaped, the tube often very long, (3)4–5(11)-lobed, the lobes mostly contorted or valvate, sometimes valvate-induplicate, rarely imbricate or quincuncial Calyx gamosepalous, the tube mostly adnate to the ovary, (3)4–5(8)-toothed or lobed, sometimes only minutely so, with open, valvate, imbricate or contorted aestivation, 1 or several lobes sometimes slightly to very considerably enlarged to form a leafy often coloured lamina Flowers rarely solitary, mostly in various terminal or axillary inflorescences, all basically cymose but variously aggregated into panicles, etc., occasionally in globose heads to an extent that the ovaries are adnate; bracts vestigial to well developed, even conspicuous; flowers usually hermaphrodite, rarely unisexual, regular or nearly so (except in Posoqueria (America)) or corolla tube rarely curved, homostylous or quite often heterostylous with 2 or rarely 3 forms (long-styled (dolichostylous), short-styled (brachystylous) or equal-styled (isostylous)) Ovary inferior, rarely half-inferior or (in Gaertnera) superior, syncarpous of 2–5 or more carpels, but predominantly of 2 and therefore predominantly 2-locular, but 3–5 or even 12 or more (eg in Urophylleae due to supplementary incomplete partitions); placentation axile or (in some Gardenieae) parietal; ovules 1-many per locule, often embedded in fleshy placentas, erect, basal or horizontal, anatropous; style simple, usually long and narrow, the “stigma” either cylindrical, clavate, or otherwise modified to form a “receptaculum pollinis” or divided into 2-many linear, spathulate or clavate lobes, the actual stigmatic surface sometimes confined to certain areas, eg the inner faces of the lobes Disc often present, 2-lobed or tubular Pollen various, mostly simple, isopolar and 3-colporate, but sometimes porate, the number of colpi or pores varying from 2–25, globose, ovoid or discoid, sometimes (in some Gardenieae) in tetrads or rarely polyads Stamens usually as many as the corolla-lobes and alternate with them, epipetalous; anthers basi-or dorsifixed, introrse, the thecae rarely multilocellate transversely Leaves opposite or occasionally verticillate, decussate, almost always entire, very rarely sparsely toothed or lobed (not in the Flora Zambesiaca area), always obviously stipulate (save in some Rubieae, where the stipules may be considered leaf-like or almost absent according to interpretation); domatia (mite associated pits or tufts of hairs) very often present axils of lateral nerves on the leaf lower surface; bacterial nodules present in a few genera Small to large trees, shrubs (including suffrutices from woody rootstocks) or less often annual or perennial herbs or woody or herbaceous climbers, sometimes spiny; raphides (tab. 1/7) present or absent; crystal sand absent or present (these two mutually exclusive) Seeds small to rather large, sometimes winged; testa cells in some tribes with very distinct pits; albumen present (save in Guettardeae); embryo straight or rarely curved, the radicle mostly longer than the cotyledons Fruit small to quite large (0|2–20 cm.), a capsule, berry or drupe or indehiscent or woody, occasionally (eg in Naulea and Morinda) united to form syncarps, (1)2-many-seeded, if capsules then loculicidal or septicidal or opening by a beak Fruit small to very large, 2–many-seeded, dehiscent (capsules either loculicidal, septicidal, opening by a beak or infrequently circumscissile, or splitting into mericarps), or indehiscent (berries, including forms with woody walls, or drupes), occasionally (eg in Nauclea and Morinda) united to form syncarps; if drupes containing one-seeded pyrenes thin- or thick-walled, opening by pre-formed germination slits or not, or less often a multi-seeded stone Ovary inferior, rarely half-inferior or superior (in Gaertnera), syncarpous of 2–10(12 or more) carpels, but predominantly of 2, and therefore predominantly 2-locular, but 3–10(12) (in Urophylleae can appear higher than actual carpel number due to supplementary incomplete partitions), or infrequently uni-locular (or partly so); ovules l–numerous per locule; placentation axile (attached to septum at apex, middle or base, usually on a placenta, often embedded in fleshy placentas) or infrequently (in some Gardenieae) parietal; ovules anatropous; style simple, usually long (but very short in Rubieae) and narrow, at the top either cylindrical, clavate, or otherwise modified to form a pollen presenter (the actual stigmatic surface sometimes confined to specific areas, eg the inner faces of the lobes**), or divided into 2–many (according to number of carpels) linear, spathulate or clavate stigmatic lobes Disk usually functioning as a nectary, positioned above the ovary inside the base of the calyx limb, annular to cushion-shaped or tubular, sometimes 2-lobed, occasionally fused to base of calyx limb-tube (Naucleeae) or absent (in wind-pollinated flowers) Pollen various, mostly simple, isopolar and 3- colporate, but sometimes porate, the number of colpi or pores varying from 2 to 25, globose, ovoid or discoid, sometimes (in some Gardenieae) in tetrads or rarely in polyads Stamens usually as many as the corolla lobes and alternate with them, epipetalous or not (some New World tribes only); anthers basi- or more often dorsifixed, introrse rarely porate (not in the Flora Zambesiaca area), the thecae rarely transversely multilocellate Corolla predominantly white but sometimes coloured, small to large and showy, gamopetalous, mostly salver- shaped to funnel-shaped or less often campanulate, occasionally tube very short, the tube rarely curved, (3)4–5(11)-lobed, the lobes contorted (mostly to the left, infrequently to the right) or valvate (sometimes induplicate or reduplicate), with or without a contact zone (i.e. thickened margin), less often imbricate (tab. 73/B–D) Calyx gamosepalous, the tube (i.e. calyx tube**) mostly adnate to the ovary, the limb with tubular part (i.e. limb-tube) present or absent, truncate or bearing minute teeth to well developed lobes (tab. 73/A1–A4), valvate (often separated by sinuses), or less often imbricate or contorted in aestivation, in a few genera 1–several lobes sometimes slightly to considerably enlarged to form a leaf-like often coloured blade (calycophyll) (tab. 73/A5) Stipules interpetiolar or occasionally intrapetiolar (mostly New World taxa); lobes shortly fused above the petioles and sometimes forming a sheath, or lobes free, entire, bifid or rarely laciniate, or particularly in herbaceous genera, fimbriate, often tipped with or separated by colleters (mucilaginous glands) (tab. 74), often with colleters inside the limb, especially towards the base Inflorescences terminal or axillary, basically thyrsoid, variously aggregated into panicles etc., occasionally in globose heads sometimes to an extent that the ovaries are adnate, or sometimes flowers solitary; bracts vestigial to well developed, even conspicuous Flowers regular or slightly to distinctly (not in Africa) zygomorphic, hermaphrodite or less often unisexual, homostylous, in which case very often protandrous, with or without pollen presentation (tab. 73/F,G), or rarely protogynous (not in Africa), or heterostylous with 2 or less often 3 forms (long-styled (dolichostylous), short-styled (brachystylous) or equal-styled (isostylous) (tab. 73/E) Small to large trees, shrubs (including suffrutices from woody rootstocks) or less often annual or perennial herbs or woody or herbaceous climbers, sometimes spiny; raphides (tab. 1/7) present or absent; crystal sand absent or present (these two mutually exclusive) Leaves opposite or occasionally verticillate, decussate, almost always entire, very rarely sparsely toothed or lobed (not in the Flora Zambesiaca area), always obviously stipulate (save in some Rubieae, where the stipules may be considered leaf-like or almost absent according to interpretation); domatia (mite associated pits or tufts of hairs) very often present axils of lateral nerves on the leaf lower surface; bacterial nodules present in a few genera Flowers regular or slightly to distinctly (not in Africa) zygomorphic, hermaphrodite or less often unisexual, homostylous, in which case very often protandrous, with or without pollen presentation (tab. 73/F,G), or rarely protogynous (not in Africa), or heterostylous with 2 or less often 3 forms (long-styled (dolichostylous), short-styled (brachystylous) or equal-styled (isostylous) (tab. 73/E) Corolla predominantly white but sometimes coloured, small to large and showy, gamopetalous, mostly salver- shaped to funnel-shaped or less often campanulate, occasionally tube very short, the tube rarely curved, (3)4–5(11)-lobed, the lobes contorted (mostly to the left, infrequently to the right) or valvate (sometimes induplicate or reduplicate), with or without a contact zone (i.e. thickened margin), less often imbricate (tab. 73/B–D) Calyx gamosepalous, the tube (i.e. calyx tube**) mostly adnate to the ovary, the limb with tubular part (i.e. limb-tube) present or absent, truncate or bearing minute teeth to well developed lobes (tab. 73/A1–A4), valvate (often separated by sinuses), or less often imbricate or contorted in aestivation, in a few genera 1–several lobes sometimes slightly to considerably enlarged to form a leaf-like often coloured blade (calycophyll) (tab. 73/A5) Pollen various, mostly simple, isopolar and 3- colporate, but sometimes porate, the number of colpi or pores varying from 2 to 25, globose, ovoid or discoid, sometimes (in some Gardenieae) in tetrads or rarely in polyads Stamens usually as many as the corolla lobes and alternate with them, epipetalous or not (some New World tribes only); anthers basi- or more often dorsifixed, introrse rarely porate (not in the Flora Zambesiaca area), the thecae rarely transversely multilocellate Ovary inferior, rarely half-inferior or superior (in Gaertnera), syncarpous of 2–10(12 or more) carpels, but predominantly of 2, and therefore predominantly 2-locular, but 3–10(12) (in Urophylleae can appear higher than actual carpel number due to supplementary incomplete partitions), or infrequently uni-locular (or partly so); ovules l–numerous per locule; placentation axile (attached to septum at apex, middle or base, usually on a placenta, often embedded in fleshy placentas) or infrequently (in some Gardenieae) parietal; ovules anatropous; style simple, usually long (but very short in Rubieae) and narrow, at the top either cylindrical, clavate, or otherwise modified to form a pollen presenter (the actual stigmatic surface sometimes confined to specific areas, eg the inner faces of the lobes**), or divided into 2–many (according to number of carpels) linear, spathulate or clavate stigmatic lobes Disk usually functioning as a nectary, positioned above the ovary inside the base of the calyx limb, annular to cushion-shaped or tubular, sometimes 2-lobed, occasionally fused to base of calyx limb-tube (Naucleeae) or absent (in wind-pollinated flowers) Seeds small to rather large, sometimes winged; testa cells in some tribes with very distinct pits; endosperm present (save in Guettardeae), entire or ruminate; embryo straight or rarely curved; radicle erect or horizontal, mostly longer than the cotyledons, orientation pendulous Fruit small to very large, 2–many-seeded, dehiscent (capsules either loculicidal, septicidal, opening by a beak or infrequently circumscissile, or splitting into mericarps), or indehiscent (berries, including forms with woody walls, or drupes), occasionally (eg in Nauclea and Morinda) united to form syncarps; if drupes containing one-seeded pyrenes thin- or thick-walled, opening by pre-formed germination slits or not, or less often a multi-seeded stone Stipules interpetiolar or occasionally intrapetiolar (mostly New World taxa); lobes shortly fused above the petioles and sometimes forming a sheath, or lobes free, entire, bifid or rarely laciniate, or particularly in herbaceous genera, fimbriate, often tipped with or separated by colleters (mucilaginous glands) (tab. 74), often with colleters inside the limb, especially towards the base Inflorescences terminal or axillary, basically thyrsoid, variously aggregated into panicles etc., occasionally in globose heads sometimes to an extent that the ovaries are adnate, or sometimes flowers solitary; bracts vestigial to well developed, even conspicuous Small to large trees, shrubs or less often annual or perennial herbs or woody or herbaceous climbers, sometimes spiny; tissues in many tribes containing abundant rhaphides Leaves opposite or verticillate, decussate, almost always entire, very rarely (not in Africa) palmatifid, toothed or finely denticulate, always obviously stipulate (save in some Rubieae, where the stipules may be considered foliar or almost absent according to interpretation), the stipules interpetiolar or intrapetiolar, entire or, particularly in herbaceous genera, variously divided into lobes or fimbriae, often tipped or separated by mucilaginous hairs known as colleters and often with colleters inside the base; the leaves in certain tribes sometimes contain small bacterial nodules Corolla small to large and showy, gamopetalous, rotate to salver-shaped or funnel-shaped, the tube often very long, (3)4–5(11)-lobed, the lobes mostly contorted or valvate, sometimes valvate-induplicate, rarely imbricate or quincuncial Calyx gamosepalous, the tube mostly adnate to the ovary, (3)4–5(8)-toothed or lobed, sometimes only minutely so, with open, valvate, imbricate or contorted aestivation, 1 or several lobes sometimes slightly to very considerably enlarged to form a leafy often coloured lamina Flowers rarely solitary, mostly in various terminal or axillary inflorescences, all basically cymose but variously aggregated into panicles, etc., occasionally in globose heads to an extent that the ovaries are adnate; bracts vestigial to well developed, even conspicuous; flowers usually hermaphrodite, rarely unisexual, regular or nearly so (except in Posoqueria (America)) or corolla tube rarely curved, homostylous or quite often heterostylous with 2 or rarely 3 forms (long-styled (dolichostylous), short-styled (brachystylous) or equal-styled (isostylous)) Ovary inferior, rarely half-inferior or (in Gaertnera) superior, syncarpous of 2–5 or more carpels, but predominantly of 2 and therefore predominantly 2-locular, but 3–5 or even 12 or more (eg in Urophylleae due to supplementary incomplete partitions); placentation axile or (in some Gardenieae) parietal; ovules 1-many per locule, often embedded in fleshy placentas, erect, basal or horizontal, anatropous; style simple, usually long and narrow, the “stigma” either cylindrical, clavate, or otherwise modified to form a “receptaculum pollinis” or divided into 2-many linear, spathulate or clavate lobes, the actual stigmatic surface sometimes confined to certain areas, eg the inner faces of the lobes Disc often present, 2-lobed or tubular Pollen various, mostly simple, isopolar and 3-colporate, but sometimes porate, the number of colpi or pores varying from 2–25, globose, ovoid or discoid, sometimes (in some Gardenieae) in tetrads or rarely polyads Stamens usually as many as the corolla-lobes and alternate with them, epipetalous; anthers basi-or dorsifixed, introrse, the thecae rarely multilocellate transversely Leaves opposite or occasionally verticillate, decussate, almost always entire, very rarely sparsely toothed or lobed (not in the Flora Zambesiaca area), always obviously stipulate (save in some Rubieae, where the stipules may be considered leaf-like or almost absent according to interpretation); domatia (mite associated pits or tufts of hairs) very often present axils of lateral nerves on the leaf lower surface; bacterial nodules present in a few genera Small to large trees, shrubs (including suffrutices from woody rootstocks) or less often annual or perennial herbs or woody or herbaceous climbers, sometimes spiny; raphides (tab. 1/7) present or absent; crystal sand absent or present (these two mutually exclusive) Seeds small to rather large, sometimes winged; testa cells in some tribes with very distinct pits; albumen present (save in Guettardeae); embryo straight or rarely curved, the radicle mostly longer than the cotyledons Fruit small to quite large (0|2–20 cm.), a capsule, berry or drupe or indehiscent or woody, occasionally (eg in Naulea and Morinda) united to form syncarps, (1)2-many-seeded, if capsules then loculicidal or septicidal or opening by a beak