Trichilia pallida Sw.

  • Authority

    Pennington, Terence D. 1981. Meliaceae. Fl. Neotrop. Monogr. 28: 1-359, 418-449, 459-470. (Published by NYBG Press)

  • Family

    Meliaceae

  • Scientific Name

    Trichilia pallida Sw.

  • Type

    Type. Swartz s.n., Hispaniola fl, fr (holotype, S).

  • Synonyms

    Portesia ovata Cav., Guarea obtusifolia Lam., Trichilia montana Kunth, Hedwigia simplicifolia Spreng., Trichilia simplicifolia, Trichilia portoricensis A.Spreng., Trichilia trinitensis A.Juss., Trichilia diversifolia A.Juss., Portesia diversifolia C.Roemer, Portesia montana C.Roemer, Portesia simplicifolia C.Roemer, Portesia trinitensis C.Roemer, Portesia echinocarpa de Vriese, Trichilia excelsa Benth., Trichilia macrophylla Benth., Trichilia echinocarpa (de Vriese) Walp., Pholacilia diversifolia (A.Juss.) Griseb., Pholacilia trinitensis (A.Juss.) Griseb., Trichilia echinocarpa (de Vriese) Walp., Trichilia weddellii C.DC., Trichilia weddellii var. parvifolia C.DC., Trichilia weddellii var. stylosa C.DC., Trichilia mollis C.DC., Trichilia riedelii C.DC., Trichilia flava C.DC., Trichilia brachystachya Klotzsch, Trichilia peruviana C.DC., Trichilia montana Kunth, Trichilia lobulata C.DC., Trichilia montana var. acutivalvis Kunth & C.DC., Trichilia laminensis Barb.Rodr., Trichilia triphylla Benth., Trichilia pauciflora Rusby, Trichilia gigantophylla Harms, Trichilia truncata Leonard, Trichilia davisii Sandwith, Trichilia subsimplex Steyerm., Trichilia skutchii P.H.Allen

  • Description

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    Species Description - Young branches puberulous, stiff pubescent, or tomentose, soon becoming glabrous, greyish to dark brown, nearly always with pustular pale small lenticels. Bud scales absent. Leaves imparipinnate, trifoliolate, or unifoliolate, (1.7-)4-15 (-55) cm long (rhachis and petiole only); petiole and rhachis terete to semiterete, usually glabrous less frequently puberulous to stiffly pubescent; petiolule 1-5 (-15) mm long or leaflets very rarely sessile. Leaflets 1-9, opposite or subopposite, usually narrowly or broadly elliptic or oblanceolate less frequently oblong or lanceolate, rarely cuneiform and then with 3-8 obscure to prominent acute teeth or lobes in upper half, apex usually attenuate, acuminate or cuspidate less frequently obtuse to truncate, base usually acute, cuneate or attenuate rarely obtuse, chartaceous to subcoriaceous, (5-)9-20(-36)[13.3] cm long, (2-)3-8(- 13.5)[5.2] cm broad, basal leaflets usually smaller, terminal leaflet often much larger than laterals, usually glabrous, or puberulous or pubescent on midrib above, lower surface usually glabrous less frequently puberulous or sparsely pubescent especially on midrib and veins, with granular red papillae, very rarely with tufts of hairs in vein axils, sparsely and irregularly glandular-striate or not; venation eucamptodromous or brochidodromous, midrib flat or slightly prominent; secondaries (6-)8-11(-14) on either side of midrib, arcuate ascending or less frequently ± straight, usually ± parallel less frequently convergent; intersecondaries short to moderate; tertiaries often oblique and parallel. Flowers unisexual, plants dioecious or rarely monoecious; inflorescence axillary or in axils of fallen leaves, sometimes cauliflorous, (0.5-)l-3(-10) cm long, fasciculate, sometimes forming a small corymbose thyrse but usually with branches spicate or irregularly cymose, densely- or laxly-flowered, puberulous to stiff pubescent, usually sessile or with a pedicel 0.5(-1) mm long above articulation, stalk below articulation 0-0.4(-1.0) cm long. Calyx usually patelliform or cyathiform rarely rotate, 1-2 mm long; calyx lobes 4(-5), ovate, triangular or attenuate often very shallow, apex acute to obtuse, 0.25-1 mm long, sparsely puberulous to densely pubescent. Petals (3-)4(-5), imbricate, (2.5-)3-5.5(-7.5) mm long, (l-)1.5-2.5(-3) mm broad, elliptic, oblong or lanceolate, apex usually acute less frequently rounded or obtuse, usually appressed puberulent outside, less frequently pubescent or subglabrous, inside glabrous or sometimes papillose. Staminal tube short cylindrical, cyathiform or urceolate, (1.5-)2-3.5(-4.5) mm long, 1.5-3 mm broad; filaments fused 1/3-3/4 their length, apex rounded, truncate or terminated by two acute appendages sometimes equalling or exceeding anthers, outside usually glabrous below and sparsely to densely long hairy above, less frequently glabrous or with a ring of hairs around base, inside usually glabrous in lower half and sparsely pubescent to densely barbate in throat, less frequently glabrous; anthers (7-)8(-10), (0.5-)0.8-1.2(-2) mm long, usually densely to sparsely hairy or glabrous; anther-odes narrower, not dehiscing, without pollen. Nectary annular or patelliform surrounding base of ovary and fused to base of staminal tube, sometimes prolonged into androecial ribs, sparsely pubescent to densely strigose rarely glabrous. Short androgynophore often present. Ovary (2-)3-(4)-locular, loculi with 2 obliquely superposed or rarely ± collateral ovules, densely stiff pubescent to strigose very rarely subglabrous; style slender, pubescent to glabrous; style-head minute, capitate. Pistil equalling or not much shorter than staminal tube; pistillode vestigial, much shorter than staminal tube, containing vestigial non-functional ovules. Capsule ovoid to globose, smooth, or obscurely to prominently verruculose or muricate, sparsely to densely puberulous or pubescent, usually with a mixture of longer stiffer hairs rarely subglabrous, (0.5-) 1-2 cm long, (2-)3(-4)-valved, valves wrinkling horizontally on drying sometimes strongly reflexed; pericarp ca. 1 mm thick; endocarp thin, cartilaginous. Usually only 1 seed developing in each fruit rarely 1-2 per valve, 0.5-1 cm long, ovoid, globose or flattened dorso-ventrally, shining; arillode fleshy, attached to seed along a thin line extending from micropyle along raphe to chalaza, growing over apex, sides, and base of seed, well developed around abortive ovules; seed coat hard. Embryo with plano-convex, collateral cotyledons; radicle apical, included or extending to surface. Exendospermous or occasionally a small amount of residual endosperm present.

  • Discussion

    1) Number of leaflets. This may vary from 1-9, and has been used as the basis of several species in the past including T. simplicifolia, T. subsimplex, T. triphylla, and T. diversifolia.

    In some areas, e.g. Veracruz, Mexico, and Amazonian Venezuela, individuals are apparently either unifoliolate or imparipinnate, and on a local basis one might be justified in regarding them as specifically distinct, although in Central America and the West Indies all intermediates between unifoliolate and imparipinnate are found. However there is a more important reason against using this character as the basis of any specific or infraspecific category, because the trend unifoliolate-trifoliolate-imparipinnate, or vice versa, appears to have arisen independently in several geographically separate areas. Evidence for this comes from the fact that in characters other than number of leaflets, the pinnate-trifoliolate-unifoliolate individuals from, for example, Amazonian Peru resemble each other more than they do plants from other geographical areas. In Amazonian Peru, there is a population with very large imparipinnate leaves and leaflets (T. gigantophylla Harms), but from the same region there have since been found individuals with large leaflets which are trifoliolate or unifoliolate. These latter also share the same leaflet venation, and characteristic capsule with prominently verrucose pericarp with the imparipinnate individuals, in a particular combination which could not be confused with that of trifoliolate or unifoliolate specimens from other areas such as the West Indies.

    On the other hand it is not possible to view these large-leaved populations from Amazonian Peru, whether they be imparipinnate, trifoliolate or unifoliolate, as a distinct taxon as they grade imperceptibly with other less distinct populations in other parts of Peru and Amazonian Brazil. In a similar way unifoliolate and trifoliolate individuals from Puerto Rico resemble closely imparipinnate individuals from the same island in a number of minor characteristics such as the rather wide angle of the secondary veins, and short petiolules.

    If the number of leaflets were used for recognising subspecies or varieties it would result in unnatural assemblages of individuals of no biological significance.

    2) Leaflet size. Apart from the example above from Amazonian Peru, individuals with large leaflets occur sporadically throughout the range of the species. In some cases they are connected to populations with normal sized leaflets by intermediates, in others they are not.

    3) Texture and indumentum of pericarp. Typically the capsule is smooth with an indumentum varying from puberulous to tomentose, and occurs throughout the range of the species. The other extreme is the prominently muricate pericarp (T. trinitensis, T. montana) in which the points are prolonged by tufts of hair. This is found from Panama, through northern and western South America, but is absent (or at least not yet collected) from Central America and southeastern Brazil, and the West Indies (except Trinidad). Between the two extremes are those individuals in which the pericarp is verrucose. These are fairly generally distributed through the range but again absent from southeastern Brazil, and the West Indies.

    Variation between the extremes is continuous, often within small areas, and is especially prevalent in western Amazonia. Variation in the surface of the pericarp is characteristic of several other species of Trichilia, e.g. T. hirta, T. moschata, and T. pseudostipularis.

    The variation in other characters of T. pallida is much more sporadic. Anthers are usually hairy, but from central Amazonia to southeast Brazil individuals often have glabrous anthers (T. weddellii, T. excelsa). Even in these areas some individuals are found with an intermediate amount or normal pubescence. The ovary is normally strigose, but in one collection from western Ecuador (Dodson 5737) it is subglabrous, although other individuals from the same population have the normal strigose ovary. In all other respects they resemble each other closely. Nectary structure shows a similar sporadic variation. In nearly all individuals it is patelliform and fused to the base of the staminal tube, but rarely it extends upwards to form androecial ribs. The use of this sort of variation to distinguish formal taxonomic units would result in a multitude of "micro" units, which would confuse rather than clarify the situation in nature.

    To summarize, the reasons for not basing subspecific or specific categories on this variation are: 1) it is too continuous, without sufficient discontinuity between the various character states either alone or in combination, 2) there is evidence that variation at least in leaflet number has followed parallel lines in several geographical areas, 3) the variation is not sufficiently correlated with geography, and 4) variation occurs among individuals of the same population as well as between individuals in separate populations. Use of the term "variety" would not distinguish between these two different biological phenomena.

    Trichilia pallida is therefore regarded as a single variable species with a series of differing populations, some intergrading in all respects, others partially distinct.

    Much of the variation today is found in regions with diverse physiography and climate such as Peruvian Amazonia, and diversification and partial isolation of populations under such conditions is to be expected. The variation cannot all be explained in terms of present day conditions, since in certain areas which are at present uniform, climatically and otherwise, we now find two or more rather distinct forms coexisting. An explanation of this phenomenon could be made in terms of past climatic changes causing contraction of the forest areas with temporary isolation of populations, and their subsequent reunification. A careful analysis of the variation and distribution of much more representative collections than are available at present might provide supporting evidence for such phenomena.

    Fruits: The mature fruit has a greenish-yellow pericarp, containing one or more black shining seeds partially surrounded by a red arillode. The arillode also develops around the aborted ovules, and the resulting mass of arillode tissue often exceeds that of the seed, so that the open fruit appears to contain a single red "seed" with only a small area of the black seed coat visible.

    Field characters: Tree to 25 m, but frequently flowering as a small treelet. Bark of smaller specimens smooth or dippled, pale grey. The flowers are greenish-cream or white, with a yellow nectary. Flowering and fruiting have been recorded in most months of the year.

    Distribution and Ecology: A variable species with a very wide distribution from Mexico through Central America, the Antilles and tropical South America as far south as northern Argentina and Paraguay. In Mexico records are confined to lowland tropical rain forest on the Gulf coast, and eastwards through Chiapas and into Peten and Alta Verapaz, Guatemala, and Belize. No records are available from Honduras or Nicaragua, but it may be expected from the Caribbean forests of these countries. The species reappears in the wet southeastern corner of Costa Rica and is then found through Panama and into Colombia. In the Caribbean it is known only from Hispaniola, Puerto Rico, and the Lesser Antilles. It is widely distributed throughout wet tropical South America, as a species of lowland and montane rain forest, ascending to 2000 m altitude in Venezuela and Colombia. In the drier areas of southeastern Brazil where the climate is seasonal, it occurs in gallery forest along rivers.

  • Common Names

    Carbón de Río, cedrillo, Pichinhuí, Siguaraya, Bois Masse, burro, caracolillo, gaita, ramoncillo, almendrillo, Obi, Pitomba, Pitombeira, Naranjuelo Cafetillo, Chivo Caspi, Uchu-Requia

  • Distribution

    Mexico North America| Chiapas Mexico North America| Veracruz Mexico North America| Guatemala Central America| San Marcos Guatemala Central America| Izabal Guatemala Central America| Petén Guatemala Central America| Alta Verapaz Guatemala Central America| Petén Guatemala Central America| Belize Central America| Toledo Belize Central America| Cayo Belize Central America| Costa Rica South America| Puntarenas Costa Rica Central America| San José Costa Rica Central America| Alajuela Costa Rica Central America| Darién Panamá Central America| Veraguas Panama Central America| Chiriquí Panamá Central America| Panamá Panama Central America| Canal Zone Panamá Central America| Haiti South America| Dominican Republic South America| Puerto Rico South America| Luquillo Puerto Rico South America| Guadeloupe South America| Dominica South America| Martinique South America| Saint Lucia South America| Colombia South America| Caquetá Colombia South America| Putumayo Colombia South America| Huila Colombia South America| Meta Colombia South America| Valle del Cauca Colombia South America| Boyacá Colombia South America| Chocó Colombia South America| Santander Colombia South America| Chocó Colombia South America| Bolivia South America| Venezuela South America| Amazonas Venezuela South America| Bolívar Venezuela South America| Barinas Venezuela South America| Lara Venezuela South America| Sucre Venezuela South America| Anzoátegui Venezuela South America| Colonia Uruguay South America| Carabobo Venezuela South America| Zulia Venezuela South America| Trinidad and Tobago South America|