Trees, shrubs, or lianas, monoecious, dioecious, or polygamous, terrestrial, amphibious, or aquatic, sometimes with erect, monopodial trunk supporting a series of horizontal, sympodial branches, armed or unarmed, not clonal; mucilaginous cells or canals often present in parenchymatous tissues; hairs various, some long, straight, sharp-pointed, eglandular, unicellular, and very thick-walled, with conic internal compartment at base (combretaceous), sometimes hairs also short-stipitate, glandular; roots occasionally with pneumatophores. Stems erect, horizontal, spreading, or prostrate; bud scales absent. Leaves persistent or deciduous, alternate and spiral, or opposite or subopposite and decussate, simple; stipules usually absent, sometimes present and minute; petiolate, petiole or base of blade often with 2 circular to elliptic and flat structures or flask-shaped nectariferous cavities; blade papery to leathery, venation pinnate, secondary veins often forming loops or smoothly arching toward margin, margins entire, surfaces glabrous or hairy; domatia often present as pits, pouches, or hair tufts. Inflorescences terminal and/or axillary, spikes or panicles [racemes]; bracts present (each flower in axil of a usually caducous bract); bracteoles 0 or 2, adnate to hypanthium. Flowers bisexual or unisexual, staminate and pistillate on same or different plants, or bisexual and staminate on different plants, or all bisexual, usually actinomorphic; perianth and androecium borne on hypan­thium; hypanthium cupulate, cylindric, or tubular, slightly to conspicuously prolonged beyond ovary and differentiated into 2 parts—proximally adnate to ovary, distally free; sepals 4 or 5, distinct or connate basally, imbricate to valvate; petals (0[4] or)5, distinct, imbricate or valvate; nectary present, usually a lobed or unlobed disc on top of ovary, often hairy; stamens [4 or]5–10[–16], inflexed in bud; filaments distinct, included to long-exserted; anthers dorsifixed, dehiscent by 2 longitudinal slits; pollen grains 3-colporate, often also with poreless furrows; pistil 2–5-carpellate, carpels connate; ovary inferior [semi-inferior], 1-locular; placentation apical; style 1; stigma 1, punctate to capitate; ovules 2–5[or 6], pendulous on elongate funiculi from apex of locule, anatropous, bitegmic, crassinucellate. Fruits drupes, ribbed and/or winged, dry to spongy or fleshy. Seed 1 per fruit, relatively large, outer portion of seed coat fibrous; embryo with usually 2 folded or spirally twisted cotyledons; endosperm absent.

Distribution

s United States, e Mexico, West Indies, Central America, South America, Asia, Africa, Australia, pantropical, warm temperate areas.

Discussion

Genera 11, species ca. 500 (5 genera, 8 species in the flora).

Combretaceae are clearly placed within Myrtales, a major angiosperm order whose mono­phyly is well supported by anatomy, embryology, and morphology (L. A. S. Johnson and B. G. Briggs 1984), as well as by plastid and nuclear DNA sequences (M. W. Chase et al. 1993; E. Conti et al. 1996, 1997; V. Savolainen et al. 2000, 2000b; D. E. Soltis et al. 2000, 2001; K. W. Hilu et al. 2003; Wang H. et al. 2009; O. Maurin et al. 2010, 2017). Morphological characteristics of Combretaceae supporting this placement include vessel elements with vestured pits, stems with internal phloem, stipules absent or present as very small lateral structures, flowers each with a short to elongate hypanthium, stamens incurved in bud, and a single style (W. S. Judd et al. 2008).

C. A. Stace (2007) recognized two subfamilies within Combretaceae: Strephonematoideae Engler & Diels (only Strephonema Hooker f. of western tropical Africa) and Combretoideae Burnett (all remaining genera); DNA-based phylogenetic analyses have supported this distinction (Tan F. X. et al. 2001, 2002; O. Maurin et al. 2010). Within the latter subfamily, two tribes are distinguished: Laguncularieae Engler & Diels (Dansiea Byrnes, Laguncularia, Lumnitzera, Macropteranthes F. Mueller) and Combreteae de Candolle [Combretum, Conocarpus, Finetia Gagnepain, Getonia Roxburgh, Guiera Adanson ex Jussieu, Terminalia]. Phylogenetic relationships within the family are imperfectly understood, and problems of generic circumscription exist (Stace).

Combretaceae are distributed pantropically, with extensions into warm temperate regions; within the geographical coverage of this flora, the species here treated are usually considered to be restricted to Florida; however, Conocarpus erectus and Laguncularia racemosa likely have become established on South Padre Island (Willacy County), Texas (B. L. Turner et al. 2003; Texas Non-Native Plants Group, http://www.texasnonnatives.org). The family is ecologically significant in coastal regions of southern and central Florida due to the presence of the mangrove species Laguncularia racemosa and the mangrove-associate C. erectus. Both occur in extensive coastal stands [along with Avicennia germinans and Rhizophora mangle]. Lumnitzera racemosa is also a mangrove species; it occurs in southern Florida only in a single naturalized population and shows potential to be an invasive species (J. W. Fourqurean et al. 2010). Combretum indicum, Terminalia catappa, T. muelleri, and, probably, T. buceras are also non-native; only the first two are considered invasive. Fruit dispersal in Conocarpus, Laguncularia, and Lumnitzera is by floating in water; the fruits of Terminalia are eaten by mammals and birds; they float and may be secondarily water-dispersed.

Combretaceae are not of great economic importance, although Terminalia buceras, T. catappa, and T. molinetii are widely used as shade trees and/or ornamentals in southern Florida due to their distinctive growth architecture, and cultivars of Conocarpus erectus with densely pubescent leaves are widely used as ornamental shrubs or small trees because of their distinctive coloration and salt tolerance. Terminalia catappa has fruits with edible kernels. Combretum indicum is often grown as an ornamental vine because of its showy flowers, which open white and change to pink and then red as the day progresses.

Selected References

None.

Lower Taxa

Combretum, Conocarpus, Laguncularia, Lumnitzera, Terminalia

Illustrations

Combretaceae Illustrations

Key

1	Leaves opposite or subopposite and decussate.	> 2
1	Leaves alternate and spiral (sometimes densely clustered distally on twigs, appearing verticillate).	> 3
2	Trees or erect shrubs; petioles with conspicuous nectar glands; leaf blades with pit-domatia abaxially toward margins at junction of secondary and lower order veins, each containing a basal gland, and with scattered salt-excreting glandular hairs; flowers incon- spicous, hypanthia free portions 1.3–2.5 mm, petals 1–1.3 mm, greenish white.	Laguncularia
2	Lianas or scandent shrubs; petioles without nectar glands; leaf blades with or without pouch-domatia abaxially, in axils of secondary and midveins, without salt-excreting glandular hairs; flowers conspicuous, hypanthia free portions 38–80 mm, petals 9–25 mm, white becoming pink then red.	Combretum
3	Stems with leaves clustered at ends of erect, short shoots, separated by longer, ± bare stem segments; trees with Terminalia-branching (branches in tiers, borne on central trunk, main axis erect and monopodial, lateral branches horizontal and sympodial).	Terminalia
3	Stems without clustered leaves on short shoots; trees or shrubs with branches ± similar, not with Terminalia-branching.	> 4
4	Leaves with pit-domatia on distal margins and apex, blade apex rounded to retuse; petioles without nectar glands; inflorescences axillary, spikes, not in spherical or oblong heads; flowers bisexual, with sepals and petals.	Lumnitzera
4	Leaves with pit-domatia abaxially at junction of veins with midveins, blade apex often acuminate to acute, occasionally obtuse, rarely rounded-mucronate; petioles with nectar glands at junction of petiole and blade base; inflorescences terminal and axillary, racemes or panicles of spherical or oblong heads; flowers unisexual (staminate and pistillate on different plants), with a single perianth whorl.	Conocarpus