Combretum fruticosum (Loefl.) Stuntz
Stace, C. A. & Alwan, A.-R A. 2010. Combretaceae. Fl. Neotrop. Monogr. 107: 1-369. (Published by NYBG Press)
Type. Venezuela. 1754-1756, Loefling s.n. (not traced). Combretum occidentale L., Syst. Nat., ed. 10, 2: 999. 1759. Nom. illegit., based on Gaura fruticosa Loefl. Combretum loeflingii Eichler in Mart., Fl. Bras. 14(2): 110. 1867. Nom. illegit., based on Gaura fruticosa Loefl. [As Exell (1953) stated, no type has been traced but the description agrees with this species, which is the most common in the region studied by Loefling.]
Combretum occidentale L., Gaura fruticosa Loefl., Combretum loeflingii Eichler, Combretum secundum Jacq., Combretum oxypetalum G.Don, Combretum micropetalum DC., Combretum tetragonum C.Presl, Combretum reticulatum C.Presl, Combretum aurantiacum Benth., Combretum warszewiczianum Eichler, Combretum benthamianum Van Heurck & Müll.Arg., Combretum gloriosum Rusby, Combretum lindbergii Rusby ex Eichler, Combretum loeflingii Eichler, Combretum farinosum var. phaenopetalum Donn.Sm., Combretum phaenopetalum (Donn.Sm.) Pittier, Combretum superbum Pittier, Combretum lepidopetalum Pittier, Combretum multidiscum Rusby, Combretum trinitense Britton, Combretum loeflingii subsp. ornithophilum Suess., Combretum wandurraganum R.E.Schult.
Species Description - Usually a woody liana to 30 m (usually much less), with stems to 18 cm diam., in the absence of support a shrub with scandent or procumbent long branches or a shrub 1-5 m or even a tree to 7 m (reports of tall trees probably refer to lianas within trees); combretaceous hairs (often very scarce) and peltate scales present. Leaves opposite, chartaceous or subcoriaceous, 3.5-19 × 1.5-10 cm, elliptic or sometimes elliptic-oblong to narrowly so, usually shortly to long acuminate but sometimes acute or obtuse at apex, cuneate to rounded or subcordate at base, often hairless or almost so but sometimes pubescent or even densely so abaxially and sparsely so adaxially, moderately to rather densely yellowish- or golden-lepidote abaxially, sparsely or very sparsely so adaxially. Venation usually eucamptodromous-brochidodromous, sometimes eucamptodromous or brochidodromous; midvein moderate, prominent; secondary veins (5-)6-9(-10) pairs, moderately spaced to distant, originating at moderately acute angles, slightly curved along length or more or less straight proximally, moderately prominent; intersecondary veins sometimes present; tertiary veins irregularly to regularly percurrent, slightly prominent; higher order veins distinct; areolation imperfect, slightly to scarcely prominent. Petiole 0.6-1.5 cm, glabrous to densely pubescent, contiguously to moderately yellow-to golden-lepidote. Inflorescences unbranched, stout, in opposite pairs in leaf-axils, 4-15 cm, usually aggregated into terminal racemes to 25(-40) cm, at the lower nodes in axils of normal leaves (usually fallen by fruiting), at the upper nodes usually without or with much reduced subtending leaves, glabrous to rather densely pubescent, densely to contiguously golden-lepidote. Flowers borne densely around rhachis but in nature all swept up to vertical position from more or less horizontal rhachis, with long exserted stamens forming bottle-brush syndrome, tetramerous, variable in size and shape, 10.5-17(-20) mm, varying from glabrous to rufous-pubescent on outside but always with the scales clearly visible, lower hypanthium 3.5-4.5(-6.5) mm, 4-angled, with pedicel-like proximal region 0.5-1.7 mm, contiguously golden-lepidote; upper hypanthium 6-12.5(-13.5) × 4-6(-12.5) mm, with narrow) infundibuliform proximal region (below disk) 1.5-4.5(-5) mm and abruptly demarcated distal deeply cupuliform region (above disk) 3-6(-7.5) mm excl. calyx lobes, moderately to densely golden-lepidote, often shortly pubescent inside even when hairless outside; calyx lobes 1.3-2.3(-3) mm, erect, acute to subacute or sometimes obtuse and apiculate, moderately to densely golden-lepidote; petals 4, 1.5-2.5(-3.2) × 0.8-1.5(-2)mm, usually falling well short of calyx lobes but sometimes reaching ca. to their apex, very rarely absent, elliptic to suborbicular, obtuse to acute, sometimes apiculate or shortly acuminate, without a basal claw, glabrous, rarely (as in type of C. lepidopetalum) sparsely lepidote abaxially; stamens 8, far exserted, with filaments 12.5-22 mm; disk densely pubescent at margin with straight or wavy hairs, with distinct free margin 0.2-1 mm; style 13-31.5 mm, exserted ca. as far as stamens, glabrous; ovules 3-6. Fruit 1.2-3 × 1-2.9 cm, oblong-elliptic or elliptic to very broadly so, rounded to truncate or slightly retuse at base, with very distinct narrow pseudostipe 0.1-0.55 cm, obtuse to rounded or retuse and usually distinctly apiculate at apex, often vinous-red, moderately to densely golden-lepidote especially on body; wings 4, 0.4-0.9 cm wide. Scales ca. 70-200 µm diam., more or less circular in outline, convexly scalloped at each marginal cell, divided entirely by radial walls or sometimes by a few to several additional tangential walls, rarely by a considerable number of additional tangential walls near the scale center; marginal cells ca. 35-70, considerably radially elongated and usually several reaching from margin to center except in cases where there are numerous tangential walls near the center. Sometimes some smaller scales (arrested development) are also present; these have many fewer walls (often all radial) with as few as 16 marginal cells.
Uses. Local uses of the lianas for baskets and mats have been noted, but the species is a popular ornamental in tropical and subtropical regions and. in heated houses, in temperate areas.Illustrations. Figs. 47b (infl), 49a-c (trichomes), 50a-e (fl), 51a & 57i-n (fr). Bernardello et al. (1994), fl, p. 297; Bot. Mag. 92: t. 5617 (1866) (as C. micropetalum); Bezerra Loloiola & Ferreira de Sales (1998), p. 178; Eichler (1867), fl, pl. 34 (as C. loeflingii); Exell (1953), fl, p. 119; Exell (1958), p. 148; Rodriguez (1993), fl, pp. 74-78; Stace & Alwan (1998), p. 343; Standley & Williams (1962), p. 274.Most of the variation in this species concerns the flowers, particularly the dimensions and pubescence, but there is a continuous range in all parameters and Exell’s (1953) concept of the species still seems fully justified. Combretum fruticosum in all its variations is rather easily recognized as it is the only “bottle-brush” species apart from C.formosum with golden scales; the flower shape, although variable, is also distinctive, there being a clear demarcation between the proximal narrowly infundibuliform and distal deeply cupuliform regions of the upper hypanthium. There are, however, some specimens that exhibit a form of gigantism, having much larger flowers than usual; their measurements have been included in the above description in parentheses. For example, Hatschbach 42241, from Parana (Brazil), has flowers 20 mm long, with the upper hypanthium 13.5 x 12.5 mm of which the cupuliform region is 7.5 mm long, and petals 3 mm long. At least one flower of this specimen has a 6-angled lower hypanthium and 6 calyx lobes, 15 stamens, and 18 ovules, but other examples with almost as large flowers seen from Rondonia and Bahia, in Brazil, and from Colombia share none or few of these irregularities. They might deserve varietal status, but on present evidence I prefer to treat them as sporadic abnormalities. On one Colombian specimen, Exell wrote that it was “near C. fruticosum but with different hypanthium and petals,” but he (1953) included it under C. fruticosum without comment. These large-flowered plants are also covered in the description by Marquete Ferreira da Silva and Valente (1996) of plants from Goiás and Tocantins (flower length, in the present sense, to 20 5 mm; her flower lengths of to 42 mm include the stamens)! she also recorded stamens to 33 mm and style to 44 mm well beyond the ranges I have encountered. For the distinctions from C. farinosum and C. formosum. under those species. see Kuntze (1898) described two varieties of C. laxum, var. viridulum and var. aurantiacum, but his “C. laxum Loefl.” refers to C. fruticosum. In fact neither variety belongs to either species; var. viridulum is C. lanceolatum, and var. aurantiacum is C. assimile (types seen at NY).Combretum farinosum var. phaenopetalum clearly belongs to C. fruticosum (type specimens at US) as also concluded by Exell (1953).Despite a note in Kew (by whom?) on three sheets of C. formosum collected by Sinclair from Tiger Island, Gulf of Fonseca, Honduras: “No doubt the type of van Heurck’s C. benthamianum in PL Nov. Hb. Heurck fasc. ii, p. 220, said to differ from Hartweg’s 526 [type of C. erianthum = C. formosum]” C. benthamianum Van Heurck & Mull. Arg. definitely belongs under C. fruticosum. Van Heurck and Muller Argoviensis (1871) stated "poils simples et ferrugineux, courts et étalés, médiocrement nombreux, qui laissent les glandes à découvert et bien visibles.” They acknowledged that Bentham had likened his C. erianthum to their plant, but pointed out that the latter differs "totalement" in various features, especially the shorter less dense calyx indumentum, which leaves the numerous scales "bien découvertes." I have also seen the types of both names. Specimens of C. fruticosum with a moderately pubescent upper hypanthium, hence precisely agreeing with C. benthamianum, are not rare (see under C. formosum for examples).None of the other synonyms represents plants with remarkable characters, except C. lepidopetalum which has scales on the petals, which is only of rare occurrence in C. fruticosum.
Distribution and Ecology: Combretum fruticosum is typically a forest liana, occupying al sorts of forests: evergreen or deciduous, tall or short, upland or lowland, dry or humid, on slopes or by rivers or lakes or in marshes, virgin, disturbed or secondary, and cut or burned. Hence it also occurs on roadsides or in scrub or sparsely wooded savannas, usually as shrubs or small trees, and is also found on the strand or among mangroves by the sea. It occurs on stony, rocky, or gravelly ground, on clays or on limestone, from sea level to 2000 m (Guatemala) but mostly from 50 to 600 m. Combretum fruticosum is the most widely distributed species of Combretaceae in the Americas, occurring from the east coast to almost the west coast of South America, and from about 23° N in western Mexico to about 33° S in Argentina and Uruguay. It was noted by Guaglianone (1998) from the provinces of Formosa, Chaco and Buenos Aires (Isla Martín García) in Argentina, but I have seen no specimens; this would extend the distribution further south (ca. 34° 12' S) than noted above, and further west than otherwise known in Argentina. It is, however, absent from the West Indies apart from Trinidad. It is only sparsely scattered in the drier parts of central Brazil and in the very humid central and lower Amazon basin, where C. lanceolatum, or C. rohrii and C. rotundifolium, respectively, are more characteristic. It is notably absent from Amapá, French Guiana, Suriname, and the adjacent parts of Guyana. In Central America it is more common than the related C. farinosum on the Atlantic side, but less common than it on the Pacific side.
Reproductive biology. The flowers are usually described as yellow, but often as red. Bernardello et al. (1994) and detailed notes on some sheets make it clear that they open greenish yellow, become bright yellow, and then fade to orange, and finally red; many specimens noted as yellow become red after drying. These comments apply mainly to the filaments, which are the main attractants, but the petals and anthers are sometimes said to change color. The color of the hypanthium is due to the indumentum: golden yellow due to scales, especially when they are full of secretion, or rufous if more pubescent. There are one or two references to white flowers. Flowering mostly October to August; fruiting through most of the year but mainly December onwards. The stiff spikes exhibit the classic bottle-brush pollination syndrome (Fig. 47) similar to that in the familiar Australian Callistemon, except that in Combretum fruticosum the spikes are aggregated into racemes and in this species the flowers are yellow. They are highly melliferous, producing a watery nectar. Although most bird-pollinated flowers are scentless, and C. fruticosum was stated as “odourless” by Bernardello et al. (1994), some labels refer to its fragrance or even an “unusual perfume.” The flowers are very attractive to birds, hummingbirds and passerines and some others, and are also regularly visited for nectar by small monkeys and butterflies. Sazima et al. (2001) reported observations by P. J. De Vries of the Pierid butterfly Phoebis philaea visiting the flowers in Costa Rica. In northern Argentina Bernardello et al. (1994) recorded, as visitors, two species of hummingbird (Chlorostilbon aureoventris and Hylocharis chrysura, Trochilidae), three species of passerine birds (Turdus rufiventris, Turdidae; Zonotrichia capensis and Molothrus badius, Emberizidae) and one butterfly (Dryas julia; Nymphalidae); 63.6%, 22.8%, and 13.6% of the total visits were by these three groups respectively. In Peru Janson et al. (1981) recorded seven species of primates as diurnal visitors: pygmy marmoset (Cebuella pygmaea), spider monkey (Ateles paniscus), tamarins (Sanguinus fuscicollis and S. imperator), squirrel monkey (Saimiri sciurus), and capuchin monkeys (Cebus albifrons and C. apella). There is a BBC wildlife film showing nectar drinking and pollen transference by a squirrel monkey and a passerine bird. Janson et al. also observed 14 species of birds as visitors in the families Trochilidae, Corvidae, Icteridae, Thraupidae, and Psittacidae. At night, some night monkeys (Aotus sp.) were seen at the flowers, but no bats. Reference by Janson et al. to earlier reports of visits by lemurs to C. fruticosum in Madagascar are based on their confusion of C. phaneropetalum Baker (the species involved, = C. albiftorum (Tul.) Jonkind) for C. phaenopetalum (Donn. Sm.) Pittier, which is indeed a synonym of C. fruticosum. Observations by Gryj et al. (1990) supposedly of this species probably refer to C. farinosum, which was included by them in their concept of C. fruticosum. Bernardello et al. (1994) found that C. fruticosum is slightly protogynous and self-incompatible and that each flower lasts for about four days, during the first three of which nectar is secreted. Pollen fertility was only about 45%, and only about 16% of flowers produced fruits.
Mafumbo, mofumbo, ciume, escova de macaco, pente de macaco, cipó-pincel, escova de macaco, pente de macaco, kadiamen, melosa, chupa-chupa, arkon nëka mukon, jagua, papamiel, peineta, cepillo, escobilla de monte, curassow comb, tietie, bejuco de escobetilla, bigote, cepillo de Moro
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