4. TERMINALIA Linnaeus, Syst. Nat. ed. 12. 2: 665, 674. 1767, name conserved.
[Latin terminus, terminal, and alis, pertaining, alluding to leaf clusters at branch tips]
Anogeissus (de Candolle) Wallich ex Guillemin & Perrottet, Fl. Seneg. Tent. 1(7): 279. 1832; Buchenavia Eichler; Bucida Linnaeus, name conserved; Pteleopsis Engler; Ramatuela Kunth; Terminaliopsis Danguy
Trees or shrubs, with Terminalia branching (branches in tiers, borne on central trunk, main axis erect and monopodial, lateral branches horizontal and sympodial), without pneumatophores. Stems erect and horizontal with leaves clustered at ends of erect, short shoots, separated by longer, ± bare stem segments. Leaves persistent or tardily deciduous (plants then briefly leafless), alternate and spiral; stipules present, appearing as fingerlike, multicellular, glandular hairs at petiole base; petiole not differentiated proximally and no part of it persistent, nectar glands absent [sometimes with sessile glands], sessile glands sometimes present at blade base; blade papery to leathery, venation brochidodromous [nearly eucamptodromous], apex acuminate, acute, mucronate, obtuse, retuse, or rounded; with or without hair-tuft or pit- [pocket-] domatia abaxially, at junction of secondary and tertiary veins with midvein, or junction of tertiary with secondary veins, sometimes also at other vein junctions, without glands. Inflorescences axillary, spikes, usually clustered on branches distally; bracteoles absent. Flowers somewhat conspicuous [inconspicuous], bisexual or bisexual and staminate on same plant, with staminate flowers distal [proximal] on spikes (or mixed with bisexual flowers); hypanthium cupulate, free portion 1–2 mm, from constriction at apex of ovary, pubescent abaxially throughout or pubescent proximally and glabrous distally [glabrous throughout]; sepals 4 or 5, pale green, yellow, or white, sometimes red-tinged, triangular to ovate-triangular, glabrous abaxially or pubescent; petals 0; stamens [4–]8–10, exserted; nectary disc lobate, atop ovary, pubescent; ovary terete, straight to slightly curved; style ± straight, free from hypanthium, pistillode absent in staminate flowers; ovules 2 or 3. Drupes radially symmetric in cross section to slightly [strongly] flattened, ovoid to ellipsoid; with 2 poorly developed ridges, 2 well-developed ridges or wings [2–5 well-developed wings], or 5 poorly developed and rounded lobes [terete and without lobes or wings]; hypanthium and calyx persistent or deciduous.
Species ca. 200 (4 in the flora): Florida; nearly worldwide in pantropics.
The flowers, with their exposed nectar glands, attract a variety of insects. Terminalia has a distinctive branching pattern, in which the main axis is erect and monopodial and the lateral branches are horizontal and sympodial. On the lateral branches, each segment originates as a lateral branch within the terminal leaf cluster of the previous segment. At first, each lateral branch grows horizontally, but eventually its apex becomes erect as a short shoot, producing a new terminal leaf cluster. Each sympodium as a whole branches where two lateral shoots continue its growth, substituting for the erect, terminal short shoot.
Terminalia buceras and T. molinetii often have been placed in Bucida, a genus traditionally separated from Terminalia by the retention of the free portion of the hypanthium and calyx on the fruit. The calyx is more or less deciduous in T. molinetii and it is persistent in three species of Madagascar and two of the Solomon Islands, which are phenetically quite different and likely not closely related to T. buceras or T. molinetii, calling this generic distinction into question (C. A. Stace 2007, 2010). A DNA-based phylogenetic analysis (O. Maurin et al. 2010) indicated that the recognition of Bucida renders Terminalia non-monophyletic. Bucida, therefore, has been placed in the synonymy of Terminalia (Stace 2007, 2010) as sect. Bucida (Linnaeus) Alwan & Stace.
SELECTED REFERENCES Fisher, J. B. and H. Honda. 1979. Branch geometry and effective leaf area: A study of Terminalia-branching pattern. 1. Theoretical trees. Amer. J. Bot. 66: 633–644. Fisher, J. B. and H. Honda. 1979b. Branch geometry and effective leaf area: A study of Terminalia-branching pattern. 2. Survey of real trees. Amer. J. Bot. 66: 645–655.