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 and without coloured juice; 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. Leaves without a persistent basal meristem.
Leaf anatomy. Stomata paracytic.
Adaxial hypodermis present (commonly). Lamina dorsiventral (mostly), or isobilateral to centric; with secretory cavities. Secretory cavities containing oil, or containing resin (yellow or otherwise brightly coloured); Secretory cavities schizogenous. Minor leaf veins without phloem transfer cells (Garcinia, Hypericum).
Stem anatomy. Young stems of 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 superficial. Nodes unilacunar. Internal phloem absent. Secondary thickening developing from a conventional cambial ring (mostly), or anomalous (? — Endodesmia). ‘Included’ phloem present (Endodesmia), or absent. Xylem with tracheids, or without tracheids; with fibre tracheids, or without fibre tracheids; with vessels. Vessel end-walls simple, or scalariform and simple. Vessels without vestured pits. Wood parenchyma apotracheal, or paratracheal.
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’; in cymes, in umbels, and in panicles. The terminal inflorescence unit 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; polysepalous, or gamosepalous (basally). 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, biochemistry. Cyanogenic, or not cyanogenic. Alkaloids present (rarely), or absent. Iridoids not detected. Arthroquinones detected (4 genera); polyacetate derived. 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). Arbutin absent. Sugars transported as sucrose (in Clusia, Garcinia). C3 and CAM. CAM recorded directly in Clusia. Anatomy non-C4 type (Calophyllum, Hypericum).
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 (1998) Eudicot; core Eudicot; Rosid; Eurosid I; 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).
Illustrations. • Hypericum. • Hypericum androsaemum. • Technical details (Hypericum, Pilosperma). • Technical details (Allanblackia). • Technical details (Clusia, Symphonia). • Technical details (Kielmeyera, Garcinia).
Quotations
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)
Cite this publication as: ‘L. Watson and M. J. Dallwitz (1992 onwards). The Families of Flowering Plants: Descriptions, Illustrations, Identification, and Information Retrieval. Version: 14th December 2000. http://biodiversity.uno.edu/delta/’. Dallwitz (1980), Dallwitz, Paine and Zurcher (1993, 1995, 2000), and Watson and Dallwitz (1991) should also be cited (see References).
