Tabebuia ochracea

(A. Gentry) A. Gentry

Corteza amarilla

(Bignoniaceae)


Sullivan, J. J., Masís, A. , Chavarría,F., Espinoza, R., Guadamuz, A. y Perez, D. 1998. Species Page de Tabebuiaochracea (Bignoniaceae), 30 Setiembre 1998. Species Home Pages, Areade Conservación Guanacaste, Costa Rica. http://www.acguanacaste.ac.cr


I. Identification
II. Geographic Distribution
III. Natural History
IV. Uses
V. How to find


I. Identification:

Trees of up to 25-30 m in height (the height of a mature dry forest canopy)and approximately 50 cm DBH (Diameter at Breast Height).

Other common names: "Cortez amarillo", or simply "cortez"or "corteza". People use these names to also refer to the otherspecies of Tabebuia in Costa Rica that produce yellow flowers (T.guayacan and T. chrysantha). The name "guayacán"can also be applied to T. ochracea (as well as T. guayacanand T. chrysantha), although they should not be confused with thereal "guayacán", Guaiacum sanctum (Zygophyllaceae).The name "guayacán" is applied to T. ochracea because,like G. sanctum, T. ochracea has extremely hard wood.

Flowers:

Tabebuia ochracea is famous throughout Central America for itsbrief, spectacular displays of large yellow flowers in the dry season. Thesedisplays last only 4-5 days and are synchronized across the trees of largeareas of forest and agroscape (see Natural History notesfor more details).

Each flower is approximately 8 cm long, and part of a terminal inflorescence(i.e., at the tip of a branch). The flowers are perfect and zygomorphic,i.e, each flower has both male and female sex organs, and it has only oneplane of symmetry. One flower lasts only one day, and an inflorescence intotal lasts only 4-5 days.

 

Fruit and seeds:

T. ochracea fruit are 20-40 cm long pods, with the two outer wallscovered in dense pale brown hairs. Inside a fruit pod are two hemispheresof tightly packed seeds separated by a central woody wall. When a fruitpod is mature and dry, the two outer walls separate to release a large numberof small, white, laterally-winged seeds (15-25 mm long). It is possiblefor fertilized flowers to develop into completely mature fruit, dispersingtheir seeds, within 3 weeks. The seeds normally disperse in the last weeksof the dry season and the first weeks of the wet season (between late Aprilto early June, with the exact weeks of dispersal varying year to year).

Leaves:

T. ochracea leaves are palmately compound, each of the 5 leafletsemerging from a central point at the end of the petiole (see photos below).Leaves are oppositely arranged, occurring in pairs along a shoot. The leavesof mature trees are characterized by dense white pubescence on the undersidesof leaves. This is especially evident in T. ochracea trees of dryforests, although less evident in trees of wetter forests. No other treespecies in the dry forests of the ACG has palmately compound leaves withpubescent white undersides.

There are marked differences between the leaves of mature trees of theforest canopy and the leaves of understory saplings. Relative to the leavesof mature trees, sapling leaves have browner hairs at much, much lower densities(see photos below). Saplings leaves are also much larger than tree leaves,with most sapling leaves 35-55 cm long while with most tree leaves are 25-35cm long. The leaflets of sapling leaves occasionally have serrations ontheir margins, a trait absent from tree leaves.

The leaves of seedlings and very young saplings (plants less than 50cm high) are often simple or palmately compound with 3 leaflets.

A leaf of a mature tree, upper surface (left) and lower surface (right):

A leaf of a sapling, upper surface (left) and lower surface (right):

[Note that these photos are not at the same scale. The saplingleaf is much larger than the tree leaf- see text.]

 

Bark and wood:

T. ochracea bark is light grey with thin shallow longitudinalfissures. The tissue below the old bark of tree limbs can be bright yellowin color. This coloration is most likely the origin of this tree species'common name, corteza amarilla (literally, "yellow bark").

There is a marked color difference between the sapwood and the heartwoodof T. ochracea wood, the outer sapwood being light tan to light greywhile the heartwood is a rich red brown. When first cut, the wood is coveredin a yellow powdery substance called "lapachol" which is releasedfrom the broken vessels. This sulfur yellow powder turns deep red in thepresence of alkaline solutions (Record and Hess 1940). The "lapachogroup" is a term that foresters apply to the woods of many differentTabebuia species throughtout latin america that produce the lapacholsubstance when cut, including T. ochracea.

 

Whole plant:

T. ochracea trees throughout most of Guanacaste are charcterizedby contorted trunks, as juvenile trees are especially adept at growing towardswherever there is most light and are also commonly attacked by the tabebuiashoot borer. This characteristic of contorted tree trunks is much less evidentin old growth forests where the trunks of T. ochracea trees can bevery straight (light sources for young trees are more vertical in old growthforests, and infestation by the tabebuia shoot borer is also less common).

The architecture of young Tabebuia rosea saplings is describedin detail by Borchert and Tomlinson (1984) and Borchert and Honda (1984),and T. ochracea sapling architecture is comparable (Borchert andTomlinson 1984, personal observation). Like T. rosea, undamaged Tabebuiasaplings are characterized by a leader branch with a vigorously growingshoot, and successive pairs of lateral branch complexes with many less vigorouslygrowing shoots. Wild saplings growing in the dry forests of the ACG arehardly ever as clean as the architectural model that Borchert and Tomlinson(1984) and Borchert and Honda (1984) describe, as the vigorously growingshoots of these leader branches are highly prone to tabebuia shoot borerdamage, and infestation over successive years quickly complicates the architectureof the sapling apex.

 

Similar species:

Tabebuia chrysantha (Jaquin) Nicholson
Tabebuia guayacan (Sekemann) Hemsley
Tabebuia impetiginosa (Martius ex De Candolle) Stadl.
Tabebuia rosea (Bertol.)DC

T. ochracea is the most common species of Tabebuia in thedry forests of the Area de Conservación Guanacaste. Two other speciesof Tabebuia are also commonly encountered in these dry forests, T.rosea and T. impetiginosa. In the ACG dry forests, only thesethree Tabebuia species have oppositely arranged palmately compoundleaves with five leaflets (all of the other species with palmately compoundleaves with five leaflets have their leaves arranged alternately). T.ochracea trees are easily distinguished from T. rosea and T.impetiginosa trees by the distinctive pubescent white undersides ofT. ochracea leaves. The leaves of T. rosea and T. impetiginosatrees lack obvious pubescence. Saplings of T. ochracea, with theirlarger, less pubescence leaves, can still be easily be distinguished fromsaplings of T. rosea and T. impetiginosa by the obviouslypubescent brown shoot walls of T. ochracea saplings. Like their treeleaves, the leaves and shoots of T. rosea and T. impetiginosasaplings are largely hairless. Of the dry forest Tabebuia species,only T. ochracea has yellow flowers.

[Note: T. rosea can be distinguished from T. impetiginosaby T. rosea's much thicker, tougher leaves and by the astringentsmell of crushed T. impetoginosa leaves that T. rosea (andT. ochracea) lack.]

T. ochracea also exists in the wetter forests of the ACG suchas in sectors Orosi and Cerro El Hacha. Two other species of Tabebuiaalso occur in the wet forests of Costa Rica, T. chrysantha andT. guayacan both of which also produce spectacular, synchronizeddisplays of yellow flowers. T. guayacan has been recently identifiedin the ACG, and it is possible that T. chrysantha also occurs there.When moving into progressively wetter forests, the leaves of T. ochraceatrees and saplings become less pubescent than their dry forest relatives,and it becomes possible to confuse T. ochracea with T. chrysantha.T. chrysantha has less dense pubescence than T. ochracea andcan have some leaves with 7 leaflets (Quesada Quesada et al. 1997). Unlikethe above species, T. guayacan leaves lack all pubescence excepton the leaf axil (with stellate trichomes) (Gentry 1980).

Nomenclature:

Tabebuia ochracea Standl. ssp. neochrysantha (A. Gentry)A. Gentry

Synonym: This species was previously named Tabebuia neochrysanthaA. Gentry.

II. GeographicDistribution:

International:

The species Tabebuia ochracea occurs naturally throughout mostof latin america, from Honduras in the north to Brazil in the south (Gentry1992). The central american subspecies T. ochracea neochrysantha thatoccurs in the ACG ranges from Honduras and El Salvador to Venezuela andTrinidad, and occurs in tropical dry forests through to the drier partsof tropical moist forests, from 0-1000 m in altitude (Gentry 1992).

In Costa Rica:

T. ochracea is common in the drier forests throughout Costa Rica,especially in the Guanacaste province. It is also common to see trees ofT. ochracea throughout the urban and agricultural areas of CostaRica, again especially in Guanacaste.

In the Area de Conservación Guanacaste:

T. ochracea can be found througout all of the drier forests ofthe ACG, occuring in parts of all of the conservation area's sectors. Itdoes not occur in the cloud forests of Cacao, Orosi, and Rincon de la Vieja,but it does occur on the drier western slopes of Rincon de la Vieja (SectorPailas) to an altitude of at least 900 m. The species occurs in the highestdensities in lowland dry forests. For example, in 1996 a 35 hectare plotof 70-80 year old dry forest in Sector Santa Rosa (called Bosque San Emilio),there were 495 T. ochracea trees greater than 3 cm DBH (Brian J.Enquist, unpublished data).

 Mapa de macro-habitats del ACG

 

III. NaturalHistory:

Summary of phenology:

DRY SEASON (late December to early May):

Trees leafless throughout the deciduous dry forest. Some plants in wetter forests and patches of evergreen old growth dry forest can maintain their leaves, but no growth occurs.

One or more mass flowering events occur mid- to late-dry season, cued by infrequent dry season cold fronts or rain spells.

Fruit pods develop to maturity in about three weeks following a flowering event.

Thousands of tiny winged seeds are released late dry season or early wet season, forming highly leptokurtic seed shadows about each parent tree.

WET SEASON (late May to early December):

Leafless trees produce new shoots and expand new leaves. Most of the year's shoot expansion occurs during the first few weeks of the wet season.

Also with in the first few weeks of the wet season, all viable seeds germinate, producing a short lived carpet of seedlings. Most soon die.

Within a month or two, tree shoot growth is terminated by apical flower primordia, which lie dormant until the following dry season.

Throughout the rest of the wet season, saplings and adult trees photosynthesize but rarely produce more leaves. There can be a short additional burst of growth in some saplings following the end of the "little dry season" in late August.

 

Life and death of seedlings:

All viable T. ochracea seeds germinate in the first month of thewet season. No further germination occurs later in the season or the followingyear.

The great majority of these seedlings do not survive the wet season,most of the surviving seedlings do not survive their first dry season, andthe growth rates of the few seedlings that survive to one year old are mostlyvery slow. For example, in July 1995, the positions of 272 first year seedlingswere mapped within 40 m of a parent tree in a 30 year old patch of SantaRosa dry forest. Of these, only 31% survived to the end of August 1995,and 3% survived through the dry season to the beginning of June 1996. By1997, all were dead. Growth rates of those seedlings that survive theirfirst years are also extremely low. In July 1995, heights were recordedfor 41 non-first year seedlings <= 20 cm high in the same patch forest.Most of the seedlings were in light shade, and some below small canopy lightgaps. Despite this, when they re-measured in July 1996, it was found thatthese plants increased in mean size from 9.9 cm (s = 3.9) to only 10.3 cm(s = 3.3), with the fastest growing individual moving from 15 cm to 20 cm(J. J. Sullivan, unpublished data). The inevitable conclusion is that thegreat majority of wild seedlings grow extremely slowly.

These observations contrast dramatically with the growth rates possiblein managed propagation that have been demonstrated at the Horizontes ExperimentalForestry station of the ACG. Seeds that are collected from trees in Januaryand February, sun dried, and potted with adequate water in light shade andwith added fertilizer and fungicide are about 40 cm high by June and readyto plant out into the field (F. Méndez Miranda, personal communication).Four year old plants in an open plantation are 1.5 m to 2 m high with manyshoots.

Growth of saplings:

Foliage and shoot growth on dry forest saplings flushes out followingthe onset of the rains in early May, with saplings leafing out over a weekbefore mature trees (Reich and Borchert 1982). Sapling shoot growth is almostentirely largely restricted to the first month after the beginning of therains, although a second flush of growth occurs in some saplings when the"little dry season" ends in August. In November-December at theonset of the dry season saplings shed their leaves (although in wetter forestand old growth dry forest this is not always the case).

T. ochracea is a shade-tolerant species requiring canopy gapsfor growth, like the related T. rosea (Brokaw 1980 (cited in Coley1983), Welden et al. 1991). The abundance of T. ochracea saplingsof various heights in the understory of Santa Rosa dry forests testifiesto their ability to survive for many years in the shade of secondary growthforests. However to grow in height enough to reach the canopy, saplingsneed to be in canopy gaps left by fallen trees. This is well illustratedby the vertical growth rates of a 1997 sample of 82 saplings between 0.5-6m in initial height in an area of 70-80 year old dry forest in Sector SantaRosa, half of which were located in the centers of treefall light gaps andhalf in neighboring understory shade . The shaded saplings grew on averageonly 3.2 cm in height (stdev = 9.1, max. = 25 cm) while the saplings inthe light grew on average 26.6 cm in height (stdev = 29.5, max. = 109 cm).That initially taller saplings did not grow in height significantly morethan the smaller saplings.

At these average 1997 growth rates, a 1 meter high saplings in a lightgap will need 107 years to reach a 30 m high dry forest canopy, while ashaded sapling will require 800 years (such a shaded sapling will be squashedby a falling branch well before it ever lives this long). This is obviouslya rough calculation, but it well illustrates the importance of canopy gapsfor the regeneration of T. ochracea.

Growth, flowering, and death of adulttrees:

At the beginning of the wet season, adult trees grow shoots and fromthe two opposite lateral buds that were below the apices of the dry season'sinflorescences and infructescences. This growth elongation halts 1-2 monthsinto the wet season when the apical meristem is terminated by the formationof apical flower primordia (Reich and Borchert 1982). These densely hairyflower primordia remain dormant until the middle of the dry season of thefollowing year. During the dry season months adult trees are leafless. Floweringoccurs synchronously across the population, cued by infrequent dry seasoncold fronts or rain spells.

The opening of flowers is synchronized both at the individual and populationscales. All of the inflorescences in the canopy of a tree open their firstflowers at the same time, 6-7 days after a day of unseasonable rain or coldair in the dry season. Flowers open at this time for most of the reproductivelymature trees in the area affected by the unseasonal weather. Thousands ofT. ochracea trees in a large portion of the Guanacaste province canbe ablaze with brilliant yellow for the same one 4-5 day period. In thedry season of some years there is more than one flowering event, and a singletree can produce a flower crop in more than one of these flowering events.

It is often claimed that T. ochraceatrees need a rain shower in the dry season to initiate their flowering.Although flowering episodes do often follow an unseasonal dry season cloudburst,this is not always the case. For example, in the 1997 dry season in SectorSanta Rosa, the April mass flowering event followed a period of cold nightswhen no rain fell (see graph). Apparentlythe trees use unseasonally cold temperatures in the dry season to cue thesynchronous opening of their flowers, whether there is associated rain ornot. They do not require a new supply of water to expand their flowers.

No self-fertilization of flowers occur within a tree canopy, with flowersdemonstrating late-acting incompatibility (Gibbs and Bianchi 1993).

Trees of T. ochracea are regularly more than 100 years old. Treedeaths are infrequent and are due to a variety of causes. Of those freshtree corpses known in Sector Santa Rosa, one had been killed by a lightningstrike, two were growing in very shallow soils and had been toppled overby the wind, one had such an unbalanced trunk that it's canopy had snappedoff, and another had its base rotted out by fungus which had entered throughan old fire scar at the base of the tree.

Flower pollinators and nectar robbers:

Many different kinds of bees, flies, butterflies, wasps, and hummingbirdscan be observed visiting T. ochracea flowers. They are pollinatedby a variety of bees, especially euglossines and anthophorids, and the nectaris commonly robbed by hummingbirds and xylocopid bees (Gentry 1983).

Herbivores, seed predators, and pathogens(see also the Tabebeuia ochracea insectherbivore list):

T. ochracea flowers are fed upon by a variety of canopy dwellingvertebrates, including spider monkeys (Ateles geoffroyi, Cebidae),howler monkeys (Alouatta palliata, Cebidae), varigated squirrels(Sciurus variegatoides, Sciuridae), and ctenosaur lizards (Ctenosaurasimilis, Iguanidae). These flowers may be a valuable source of waterat the end of the dry season for these animals. Fallen flowers are commonlycollected by leaf-cutter ants (Atta cephalotus, Formicidae).

Developing T. ochracea fruits are fed upon by white-faced monkeys(Cebus capuchinus, Cebidae) which rub off the hairs, open up fruitsand eat out the young seeds (Lisa Rose and Katherine MacKinnon, personalcommunication). The developing seeds are also fed upon by a species of caterpillar(Pyralidae), which mine through the seeds. The caterpillars can destroythe great majority of the developing seeds in heavily infested fruit pods.

Above ground vegetative tissues of T. ochracea in Santa Rosa arefed on by at least XX species of insect but no vertebrates (for more details,see Tabebuia ochracea insect herbivorelist). Most of these insect species feed on fully expanded leaves ofunderstory plants (personal observation). Shoots are destroyed by the tabebuiashoot borer (Lepidoptera: Pyralidae). A coreid and a pentatomid bug bothsuck from the vascular tissue of young growing shoots, and the scarabidbeetle Euphoria limatula scrapes notches out of the sides of youngshoots at the beginning of the wet season. Monochroa sigimarini caterpillars(Lepidoptera: Gelechiidae) feed on the expanding young leaves (occasionallyalso damaging the shoot tip), as do a species of unidentified fly larvawhose feeding distorts leaf development. Young seedlings have small, simpleor trifoliate leaves that are ignored by most of the T. ochraceaherbivores, although early instar larvae of Ecpatheria icasia (Lepidoptera:Arctiidae) have been observed feeding on small seedlings, and leaf cutterants (Atta cephalotus, Formicidae) often remove a large proportionof the cotyledons of newly germinated seeds. In the adult crown I have observedMonochroa larvae (when new leaves flush at the beginning of the wetseason), Euphoria limatula, the chrysomelid alticin beetle Megistopsnr. costaricensis, and infrequently the tabebuia shoot borer.

IV. Uses:

Bark:

The anti-cancer properties of bark extracts from several Brazilian Tabebuiaspecies, including Tabebuia impetiginosa, have produced muchinterest in the bark chemistry of the genus both among biochemists (e.g.,Kang et al. 1990, Blatt et al. 1996, Pinto et al. 1997) and the alternativemedicine industry (e.g., see http://alternativemedicine.com/issue7/pharm.html),and includes T. ochracea (e.g., Zani et al. 1991). In the Andes,the inner bark of T. impetiginosa is traditionally scraped and madeinto a tea with reported medicinal propertes, and the American Health andHerbs company which now sells T. impetiginosa at $13.10 for 2 OZ.claims that the extract can be used for everything from skin sores and wartsto asthma to arthritis to snake bites to killing viruses (including AIDS!)(http://www.cyberline.com/herbs/singles.htm, 1996). Drink with a grain ofsalt!

Wood:

The lapacho Tabebuia woods, including T. ochracea, arerenowned for their exceptional strength and durability, being perhaps thehardest, heaviest, most durable woods of any group of neotropical trees(Gentry 1980). The heartwood is very reistant to attack by fungi and termites(Chudnoff 1984). The classic example of this are beams of T. guayacan,another lapacho Tabebuia, that were used to build the cathedral ofOld Panama city and are still perfectly sound after exposure to weathersince the destruction of the city over 300 years ago (Record and Hess 1940).

These hard, heavy, and durable qualities make lapcho Tabebuia woodsvaluable for many uses. The woods are especially used for the constructionof outdoor furniture, railroad crossties, heavy construction, tool handles,industrial flooring, etc. (Chudnoff 1984, San Román et al. 1981,Carpio Malavassi 1995). Tabebuia wood of the lapacho group (whichalso commercially goes by the name "ipe") was one of the maintimbers used by outdoor furniture manufacturers in the USA (now mostly replacedby plantation teak), and is now used in the US for flooring (and is apparentlyspecified for US Army truck beds) (John Curtis, personal communication).In Costa Rica, these woods were traditionally used to make the parts ofox-pulled carts, and are now one of the woods favored for the beds of convertedpickup trucks. The abrupt difference in color between the sapwood and heartwood,combined with the wood's "magnificent finish" (San Románet al. 1981) also make it useful for furniture, household objects, touristgift items, and decorative veneers (Gentry 1980, Chudnoff, 1984). The timberis further useful as a pulp for paper (Carpio Malavassi 1995, San Románet al. 1981).

T. ochracea is one of numerous native Costa Rican trees beingscreened by the Horizontes Experimental Forestry Station of the ACG forpotential use in reforestation and plantation forestry projects. Despitehaving "one of the finest woods in Costa Rica" (Nichols et al.1991 p. 21, translated), the development of T. ochracea as a highquality timber species is largely prevented by its very slow growth andoften twisted form when growing in pastures and young forests (Nichols etal. 1991, Edgar Viquez (MACORI, Liberia), personal communication).

Whole plant:

T. ochracea trees are highly valued as ornamentals due to theirspectacular dry season shows of flowers, and can commonly be found plantedin towns and cities, within and along the borders of farms, and occasionallyfor use as living fence posts. Seeds of T. ochracea can even be purchasedin San Jose tourist gift stores (under the name "Golden Goddess").

V. How to find:

Tabebuia ochracea is a common tree of the deciduous dry forestsof Sector Santa Rosa, as well as being common in the drier parts of manyother ACG sectors. There are several large saplings growing within the SantaRosa administration area (for example, outside the computer room of theCentro de Investigación) and in the forests surrounding this area(for example, along Sendero Lupo- the plant identification trail). For comparision,there are saplings of T. rosea and T. impetiginosa growingoutside of the Programa de Educación Biologica (PEB) building inthe Santa Rosa administration area. T. ochracea is also common alongthe sides of the panamerican highway between Liberia and Sector Santa Rosa.

To find a large number of T. ochracea trees, it is best to searchduring a mass flowering event (usually 4-5 days of March or April, withthe exact dates varying between years and areas). At this time, adult treesare visually advertising their locations to pollinators and a large numberof trees can be quickly located from any any high site with a view.


Acknowledgements:

The ACG Plant Species Home Pages an Species Pages project was made possibleby the Area de Conservación Guanacaste/CR, with assistance from INBio(CR), ICBG of the Foggarty Center, the NIH (USA), and the NSF (USA).


Cited Literature:

Blatt, C. T., Salatino, A., and Salatino, M. L. F. 1996. Flavonoids ofTabebuia caraiba (Bignoniaceae). Biochemical Systematics & Ecology,24:89.

Borchert, R. and Honda, H. 1984. Control of development in the bifurcatingbranch system of Tabebuia rosea: a computer simulation. Botanicalgazette, 145:184-195.

Borchert, R. and Tomlinson, P. B. 1984. Architecture and crown geometryin Tabebuia rosea (Bignonaceae). American Journal of Botany, 71:958-969.

Carpio Malavassi, I. A., 1995. Maderas de Costa Rica: 150 especies forestales.Editorial de la Universidad de Costa Rica, Costa Rica.

Chudnoff, M. 1984. Tropical timbers of the world. Agricricultural. Handbook607: U.S. Department of Agriculture, Forest Service, Washington, DC.466p. [see text extract at http://wissago.uwex.edu/test/fpl-save/ttw/5-1.html]

Gentry, A. H. 1980. Bignoniaceae- Part 1 (Crescentieae and Tourrettieae).Flora Neotropica Monograph Number 25. The New York Botanical Garden, NewYork.

Gentry, A. H. 1983. Tabebuia ochracea spp. neochrysantha (Guayacán,Corteza, Cortes, Corteza amarilla). pp. 335-336 in D. H. Janzen (ed.), CostaRican Natural History. University of Chicago Press, Chicago.

Gentry, A. H. 1992. Flora Neotropica: Bignoniaceae-Part II (Tribe Tecomeae).Flora Neotropica Monograph Number 25 (II), The New York Botanical Garden,New York.

Gibbs, P. E. and Bianchi, M. 1993. Post-pollination events in speciesof Chorisia (Bombacaceae) and Tabebuia (Bignoniaceae) withlate-acting self-incompatibility. Botanica Acta, 106:64-71.

Kang, W. B., Sekiya, T., Toru, T., and Ueno, Y. 1990. A new route tonaphtho(2,3-b)furan-4,9-diones from thio-substituted 1,4-naphthoquinones.Journal of the Chemical Society Perkin Transactions I , 0(3): 441-446.

Nichols, D. and González, E. (eds.) 1991. Especies nativas y exóticaspara la reforestación an la zona sur de Costa Rica. Memoria de IIEncuentro sobre Especies Forestales. 12-14 de febrero de 1991, JardínBotánico Wilson, San Vito de Costa Brus, Costa Rica. UniversidadEstatal a Distancia (UNED) / Organización para Estudios Tropicales(OET) / Dirección General Forestal (DGF), Costa Rica.

Pinto, A. V., Pinto, C. N., Pinto, M. D. C. F. R., Rita, R. S., Pezzella,C. A. C., and De Castro, S. L. 1997. Trypanocidal activity of syntheticheterocyclic derivatives of active quinones from Tabebuia sp.. Arzneimittel-Forschung,47:74-79.

Quesada Quesada, F. J., Jiménez Madrigal, Q., Zamora Villalobos,N., Aguilar Fernández, R., and González Ramírez J.1997. Arboles de la Peninsula de Osa. Instituto Nacional de Biodiversidad,Costa Rica.

Record, S. J. and Hess, R. W. 1940. American timbers of the family Bignoniaceae.Tropical Woods, 63:9-37.

Reich, P. B. and Borchert, R. 1982. Phenology and ecophysiology of thetropical tree, Tabebuia neochrysantha (Bignonaceae). Ecology, 63:294-299.

San Román, M., González T., G., López, A., RiveraGonzález, D., Méndez Salazar, L., Bonilla S., L. Ma., CarpioMalavassi B., I. Ma., and Muñoz, A. 1981. Propiedades y usos de cuarentay ocho especies maderables de llanos del cortes, Guanacaste. Laboratoriode Productos Forestales, CATIE-UCR-MAG, Universidad de Costa Rica.

Zani, C. L., De Oliveira, A. B., and De Oliveira, G. G. 1991. Furanonapthoquinonesfrom Tabebuia ochracea. Phytochemistry, 30(7): 2379-2381.


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