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The families of flowering plants

L. Watson and M. J. Dallwitz

Guttiferae Juss.

Alternatively Clusiaceae Lindl. (nom. altern.).

Including Calophyllaceae J.G. Agardh, Cambogiaceae Horan., Garciniaceae Dum, Hypericaceae Juss., Moronobeaceae Miers, Polyadelphaceae Dulac, Symphoniaceae Presl; excluding Bonnetiaceae.

Habit and leaf form. Trees, shrubs, herbs, and lianas; with coloured juice (this resinous), or non-laticiferous; bearing essential oils, or without essential oils; resinous, or not resinous. Self supporting, or epiphytic, or climbing. Mesophytic. Leaves opposite, or whorled; ‘herbaceous’, or leathery; petiolate to sessile; gland-dotted (commonly, conspicuously), or not gland-dotted; without marked odour; simple; epulvinate. Lamina entire; pinnately veined; cross-venulate. Leaves exstipulate. Lamina margins entire. Leaf development not ‘graminaceous’.

Leaf anatomy. The leaf lamina dorsiventral (mostly), or bifacial to centric. Stomata nearly alwys mainly confined to one surface; paracytic (commonly), or anomocytic, or cyclocytic, or actinocytic (being especially variable in Hypericum, sometimes on the same leaf). Hairs present (but seldom numerous), or absent; when present, eglandular; unicellular, or multicellular. Unicellular hairs branched (rarely), or simple (see illustration). Multicellular hairs when found, uniseriate. Complex hairs rarely present; when found, stellate. Adaxial hypodermis present (commonly), or absent. Lamina universally with secretory cavities (visisble as opaque or translucent dots). Secretory cavities containing oil, or containing resin (yellow or otherwise brightly coloured); schizogenous. The mesophyll commonly containing crystals. The crystals druses, or solitary-prismatic. Minor leaf veins without phloem transfer cells (Garcinia, Hypericum).

Axial (stem, wood) anatomy. Young stems of the herbs cylindrical, or tetragonal, or oval in section (often ridged). Secretory cavities present; with resin, or with oil. Cork cambium present; initially deep-seated, or initially superficial. Nodes unilacunar. Primary vascular tissues comprising a ring of bundles; collateral. Internal phloem absent. Cortical bundles absent. Medullary bundles absent. Secondary thickening developing from a conventional cambial ring (mostly), or anomalous (? — of the foraminate type, reported in Endodesmia). Primary medullary rays narrow.

The wood diffuse porous (to no more than slightly ring porous in Hypericum). The vessels small to medium; solitary, or radially paired, or in radial multiples, or in tangential arcs. The vessel end-walls simple, or scalariform and simple. The vessels without vestured pits. The axial xylem with tracheids (usually), or without tracheids (?); very commonly with vasicentric tracheids, or without vasicentric tracheids; with fibre tracheids, or without fibre tracheids; with libriform fibres, or without libriform fibres; including septate fibres (rarely), or without septate fibres. The fibres without spiral thickening. The parenchyma when present, apotracheal (especially in Hypericum), or paratracheal. The secondary phloem not stratified. ‘Included’ phloem present (Endodesmia), or absent. The wood not storied.

Reproductive type, pollination. Plants hermaphrodite (Hypericaceae s. str.), or monoecious, or andromonoecious, or gynomonoecious, or dioecious, or polygamomonoecious.

Inflorescence, floral, fruit and seed morphology. Flowers solitary (rarely), or aggregated in ‘inflorescences’; when aggregated, in cymes, or in umbels, or in panicles. The ultimate inflorescence units cymose. Inflorescences terminal; cymose, often umbellate or paniculate. Flowers bracteolate (the two bracteoles often close up under the calyx and not clearly distinguishable from it), or ebracteolate; medium-sized, or large; regular; cyclic, or partially acyclic (often partially spiral). Commonly the perianth acyclic, or the androecium acyclic, or the perianth acyclic and the androecium acyclic. Free hypanthium absent. Hypogynous disk absent, or present; of separate members.

Perianth with distinct calyx and corolla, or sequentially intergrading from sepals to petals; 4–12(–20); free; 2 whorled (if whorled). Calyx generally 2–6; 1 whorled; basally gamosepalous, or polysepalous. Calyx lobes markedly longer than the tube. Calyx regular; imbricate. Corolla 2–6 (-14); 1 whorled; polypetalous, or gamopetalous (sometimes basally connate). Corolla lobes markedly shorter than the tube. Corolla imbricate, or contorted; regular; yellow, or white. Petals clawed, or sessile.

Androecium 3–4 (rarely), or 20–100 (i.e. usually ‘many’). Androecial members branched (usually, apparently), or unbranched; when many (i.e. usually), maturing centrifugally (the members individually, those within bundles, and the bundles themselves); free of the perianth and adnate, or free of the perianth; free of one another, or coherent (often grouped into bundles, sometimes united into a tube or even united at their apices); 2–5 adelphous (when in separate bundles), or 1 adelphous (when A united into a tube); 1 whorled, or 2 whorled, or 3 whorled (or spiralled). The androecial bundles when bundled, opposite the corolla members (and often adnate to them). Androecium exclusively of fertile stamens, or including staminodes. Staminodes when present, 2–50 (i.e. few to ‘many’); external to the fertile stamens (commonly the outer members staminodal). Stamens (3–)5–100 (usually ‘many’); diplostemonous (rarely), or triplostemonous to polystemonous (usually); when bundled, alternisepalous. Anthers separate from one another (usually), or cohering (occasionally); dehiscing via longitudinal slits; introrse (usually), or extrorse (rarely); bisporangiate. Endothecium developing fibrous thickenings. Anther epidermis persistent. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or isobilateral, or decussate. Anther wall initially with one middle layer, or initially with more than one middle layer. Pollen shed in aggregates (e.g. Kielmeyera), or shed as single grains; when aggregated, in tetrads. Pollen grains aperturate; (2–)3(–5) aperturate; colporate; 2-celled.

Gynoecium (1–)3 carpelled, or 5(–13) carpelled (or more). The pistil 1 celled, or 3–13 celled. Gynoecium syncarpous; synovarious, or synstylovarious, or eu-syncarpous; superior. Ovary 1(–3) locular, or 5(–13) locular (as many locules as G, or unilocular by intruded placentae failing to reach the middle). Gynoecium stylate, or non-stylate. Styles when present, 1, or 3, or 5(–13); partially joined; attenuate from the ovary; apical. Stigmas 1, or 3, or 5(–13); sometimes peltate; wet type; non-papillate; Group IV type. Placentation when ovary unilocular (i.e. rarely), parietal (on intruded placentae); usually axile. Ovules (1–)2–50 per locule (i.e., to ‘many’); horizontal, or ascending; arillate (often), or non-arillate (Hypericaceae s. str.); anatropous, or hemianatropous; bitegmic; tenuinucellate. Outer integument contributing to the micropyle. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; proliferating (up to 7 cells, in Hypericum), or not proliferating; ephemeral, or persistent. Synergids pear-shaped, or hooked (sometimes with filiform apparatus). Endosperm formation nuclear. Embryogeny solanad.

Fruit fleshy, or non-fleshy; dehiscent, or indehiscent; a capsule, or a berry, or a drupe. Capsules septicidal, or septicidal and loculicidal (Eliaea). Seeds non-endospermic; winged, or wingless. Embryo rudimentary at the time of seed release to well differentiated. Cotyledons 2 (sometimes reduced). Embryo chlorophyllous (2/6); straight to curved.

Seedling. Germination phanerocotylar, or cryptocotylar.

Physiology, phytochemistry. C3 and CAM. CAM recorded directly in Clusia. Anatomy non-C4 type (Calophyllum, Hypericum). Sugars transported as sucrose (in Clusia, Garcinia). Cyanogenic, or not cyanogenic. Alkaloids present (rarely), or absent. Anthraquinones detected (4 genera); polyacetate derived. Arbutin absent. Iridoids not detected. Proanthocyanidins present; cyanidin, or cyanidin and delphinidin. Flavonols present, or absent; quercetin, or kaempferol and quercetin, or quercetin and myricetin. Ellagic acid absent (6 species, 3 genera).

Geography, cytology. Temperate to tropical. Cosmopolitan. X = 7, 8, 9, 10.

Taxonomy. Subclass Dicotyledonae; Crassinucelli (? — tenuinucellate, but polypetalous, bitegmic ovules, etc.). Dahlgren’s Superorder Theiflorae; Theales. Cronquist’s Subclass Dilleniidae; Theales. APG 3 core angiosperms; core eudicot; Superorder Rosanae; fabid; Order Malpighiales.

Species 1000. Genera about 40; Allanblackia, Calophyllum, Caraipa, Chrysochlamys, Clusia, Clusiella, Cratoxylon, Dystovomita, Eliaea (Eliea), Endodesmia, Garcinia, Haploclathra, Harungana, Havetia, Havetiopsis, Hypericum, Kayea, Kielmeyera, Lebrunia, Lorostemon, Mahurea, Mammea, Marila, Mesua, Montrouziera, Moronobea, Neotatea, Oedematopus, Pentadesma, Pilosperma, Platonia, Poeciloneuron, Psorospermum, Quapoya, Santomasia, Symphonia, Thornea, Thysanostemon, Tovomita, Tovomitidium, Triadenum, Vismia.

Economic uses, etc. Edible fruit from Garcinia (mangosteen), Mammea (mammee apple, mamey).


So then about her brow
They bound Hypericum, whose potent leaves
Have sovereign power o’er all the sullen fits
And cheerless fancies that besiege the mind
(Alfred Lear Huxford, quoted by Ann Pratt, ‘Wild Flowers’ (1857) - Hypericum perforatum)

Illustrations. • Technical details: Hypericum. • Technical details: Allanblackia (Thonner). • Technical details: Clusia, Chrysopia (= Symphonia). • Technical details: Garcinia, Pilosperma. • Technical details: Kielmeyera, Garcinia (Lindley). • Hypericum androsaemum, A. x inodorum (as H. elatum) and H. hircinum: Eng. Bot. 264–266, 1864. • Hypericum calycinum, H. perforatum and H. maculatum (as H. dubium): Eng. Bot. 267–269, 1864. • Hypericum undulatum (as H. boeticum), H. linariifolium, H. pulchrum: Eng. Bot. 270bis, 272, 273 (1864). • Hypericum montanum and H. elodes: Eng. Bot. 275 and 276, 1864. • Hypericum androsaemum, A. perforatum A. pulchrum (B. Ent. compilation, 1824–35). • Calophyllum spp.: leaf hairs (Solereder, 1908). • Calophyllum, Clusia, Garcinia, Oedematopus: leaf anatomical details (Solereder, 1908). • leaf anatomical details of Caraipa, with Theaceae (Cleyera and Freziera) and Actinidiaceae (Saurauia): Solereder, 1908.

