Taxonomic Revisions in the Polyphyletic Genus Tabebuia s. l. (Bignoniaceae)

SUSAN O. GROSE1 and R. G. OLMSTEAD

Department of Biology, University of Washington, Box 355325, Seattle, Washington 98195 U.S.A.1Author for correspondence (sheydler@smith.alumnae.net)

Communicating Editor: James F. Smith

ABSTRACT. Recent molecular studies have shown Tabebuia to be polyphyletic, thus necessitating taxonomicrevision. These revisions are made here by resurrecting two genera to contain segregate clades of Tabebuia.Roseodendron Miranda consists of the two species with spathaceous calices of similar texture to the corolla.Handroanthus Mattos comprises the principally yellow flowered species with an indumentum of hairs covering theleaves and calyx. The species of Handroanthus are also characterized by having extremely dense wood containingcopious quantities of lapachol. Tabebuia is restricted to those species with white to red or rarely yellow flowers andhaving an indumentum of stalked or sessile lepidote scales. The following new combinations are published:Handroanthus arianeae (A. H. Gentry) S. Grose, H. billbergii (Bur. & K. Schum). S. Grose subsp. billbergii, H.billbergii subsp. ampla (A. H. Gentry) S. Grose, H. botelhensis (A. H. Gentry) S. Grose, H. bureavii (Sandwith) S.Grose, H. catarinensis (A. H. Gentry) S. Grose, H. chrysanthus (Jacq.) S. Grose subsp. chrysanthus, H. chrysanthussubsp. meridionalis (A. H. Gentry) S. Grose, H. chrysanthus subsp. pluvicolus (A. H. Gentry) S. Grose, H. coralibe(Standl.) S. Grose, H. cristatus (A. H. Gentry) S. Grose, H. guayacan (Seemann) S. Grose, H. incanus (A. H. Gentry)S. Grose, H. lapacho (K. Schum.) S. Grose, H. pulcherrimus (Sandwith) S. Grose, H. pumilus (A. H. Gentry) S. Grose,H. riodocensis (A. H. Gentry) S. Grose, H. selachidentatus (A. H. Gentry) S. Grose, H. serratifolius (Vahl) S. Grose,H. spongiosus (Rizzini) S. Grose, H. subtilis (Sprague & Sandwith) S. Grose and H. uleanus (Kraenzl.) S. Grose.

KEYWORDS: Handroanthus, Roseodendron, Tabebuia, taxonomy.

Tabebuia, as currently circumscribed with 100species, is the largest genus in Bignoniaceae and isdistributed from the southwestern U.S. to northernArgentina and Chile. Over the course of itstaxonomic history, it has been split and re-assembled several times, as researchers interpretedthe morphological diversity in different ways. Thewide range of morphological diversity suggeststhere may be more than one lineage includedwithin the traditional concept of Tabebuia. Recentstudies (Spangler and Olmstead 1999; Grose andOlmstead 2007) have confirmed that Tabebuia, ascurrently delimited, is polyphyletic. Therefore, thegeneric boundaries need to be redefined inTabebuia and related genera. The goal of this paperis to revise the generic classification to be consis-tent with its phylogeny.

The name Tabebuia has a long and convolutedhistory (Gentry 1969). This name was first publishedby de Candolle (Candolle 1838) who applied it tobignoniaceous trees with ‘‘simple’’ leaves. Howev-er, the concept of Tabebuia came to be expanded toencompass a large amount of morphological di-versity. As researchers examined and monographedthis taxon, a number of concepts of Tabebuiaemerged, producing a labyrinthine synomomy(Candolle 1838; Raffinsque 1838; Sprague andSandwith 1932). In addition, there was confusionas to the boundaries between Tecoma and Tabebuia(Miers 1863; Seeman 1863; Bureau 1864; Schumann1894; Rheder 1913). Much of the confusion betweenTecoma and Tabebuia was sorted out by Britton(Britton 1915). However, disagreement as to the

delimitation of Tabebuia has continued into recenttimes (Mattos 1970; Gentry 1972).

Mattos (1970) divided Tabebuia into two differentgroups, indicating that a group of Brazilian taxaknown as ‘‘ipes’’ should not remain in Tabebuia.His reasoning appears to have been based on theoriginal concept for Tabebuia, which he defined ashaving simple leaves and an ovary that in crosssection has 3–4 ovules/locule. This morphology isconsistent with the type of Tabebuia, T. cassinoides,but is the most exclusive concept for Tabebuia sinceit was proposed by de Candolle (1838). Neither didMattos think the ipes belonged in Tecoma, due totheir palmately compound leaves. He created thegenus Handroanthus for those species with pal-mately compound leaves and 8–9 series of ovules/locule, and typified this genus with Handroanthusalbus (Tabebuia alba).

Gentry (1972, 1992) included Handroanthus inTabebuia, insisting that Tabebuia as previously de-lineated is natural. Gentry (1972) was adamant thatthe large genus Tabebuia should not be broken upfurther, and expressed his ‘‘sincere hope that futurestudents of Bignoniaceae will consider the case ofHandroanthus before succumbing to further parox-ysms of unwarranted splitting’’ (Gentry 1972).

Gentry (1992) recognized some groups withinTabebuia, however. In his treatment for FloraNeotropica he defined 10 species groups althoughhe placed two species, T. arimaoensis and T.heterophylla, in both groups 9 and 10. The largestof Gentry’s groups are primarily species endemicto the Greater Antilles (55 species). He did not use

Systematic Botany (2007), 32(3): pp. 660–670# Copyright 2007 by the American Society of Plant Taxonomists

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phylogenetic terminology to describe the relation-ships, but he clearly believed that this group ofAnillean species was monophyletic and sister toa group of mainland species that are spreadthroughout the continental part of the genus’ range(Gentry 1992).

Tabebuia donnell-smithii was placed in Tabebuia byRose (1892) with some reservation: ‘‘The species,while not agreeing in all respects with Tabebuia,answers better to this than to any other knowngenus. In its inflorescence and ribbed pods it ismore like Godmania or Cybistax but does not agreein other particulars.’’ This species was later movedto Cybistax by Seibert (1940). However, Miranda(1965) transferred this species to a new genus,Roseodendron, segregating it from Cybistax on thebasis of calyx texture and shape, sessile nature ofits ovary, and indumentum of branched hairs.

Grose and Olmstead (2007) conducted a molecu-lar phylogenetic study of the Tabebuia Alliance,those Neotropical Bignoniaceae with palmately-compound leaves. That study included 25% of allTabebuia species and had representatives from all10 of Gentry’s (1992) species groups. The findingsshowed that Tabebuia consists of three clades.‘‘Tabebuia goup I’’ is sister to the Caribbeanendemic Ekmanianthe. ‘‘Tabebuia group II’’ is sisterto Crescentieae+Spirotecoma. The third clade isrepresented by T. donnell-smithii and is sister toTabebuia group II +Crescentieae+ Spirotecoma.

MATERIALS AND METHODS

Characters to support the phylogeny of Grose andOlmstead (2007) are based on herbarium holdings at US,MO and NY and published morphological and anatomicalstudies (Gentry 1992; dos Santos and Miller 1992). Herbariumspecimens were consulted for morphological characters(Appendix 1). Leaf surfaces and calices were examined witha scanning electron microscope (SEM). Specimens wereprepared for the SEM according to the protocol establishedin Matthews and Endress (2004)

MORPHOLOGICAL CHARACTERS DEFINING CLADES

Each of the three clades of Tabebuia: Tabebuiagroups I and II and T. donnell-smithii is supportedby several morphological characters. Gentry de-scribed the importance of the calyx in Bignoniaceaetaxonomy especially in tribe Bignonieae (Gentry1980). This study confirms previous observationsthat calyx morphology as well as indumentum,wood anatomy, flower color, and fruit morphologyhave been important in defining the three groupsof Tabebuia in this clade (dos Santos and Miller1992).

Tabebuia group I shares the characters of anindumentum of lepidote scales and a spathaceous,sometimes irregularly bi- or trilabiate calyx

(Fig. 1C, F). The lepidote scales in a group ofspecies (such as Tabebuia pilosa) may be stalked.The wood anatomy of Tabebuia group I is distinc-tive; it is lightweight, with medium specific gravityof 0.4–0.74, and lacks lapachol (dos Santos andMiller 1992). The heartwood is indistinct from thesapwood and the rays are 1–2 cells in width (dosSantos and Miller 1992). This group has leaves are‘simple’ (or unifoliolate) or 3–7(9)-merous, depend-ing on species (Gentry 1992). The fruits are linear-cylindrical and smooth on the surface (Gentry1992). The flowers of Tabebuia group I are white tored in color, and often have a yellow throat (Fig 1I).Two species belonging to this clade, Tabebuia aureaand the unsampled Tabebuia nodosa have yellowflowers but otherwise share the above character-istics.

Tabebuia group II can be distinguished by havingan indumentum of simple to stellate or dendroidhairs (Fig. 1A, D). Lepidote scales are also usuallypresent, but may be obscured by the hairs. Thecalyx is campanulate to cupular and 5-dentate(Fig. 1D). The wood is among the heaviest andhardest known, with a basic specific gravitygreater than 0.74 (dos Santos and Miller 1992). Itis also distinctive among Tabebuia s. l., in havingvery large pits (ranging from 7–14 mm) in theirvessel elements (dos Santos and Miller 1992). Onlytwo species in this group, T. pumila and T.selachidentata, are described as occasionally havingunifoliolate leaves (Gentry 1992). In all other cases,the leaves are digitately 3–9 foliolate. The fruits inthis group are linear and slightly costate to smooth,and often are densely tomentose. The flowers inTabebuia group II are generally yellow (Fig. 1G) or,in four species, T. selachidentata, T. barbata, T.heptaphylla and T. impetiginosa, magenta withyellow throat (Gentry 1992).

Tabebuia donnell-smithii represents a group dis-tinguished by its unique calyx (Fig. 1E). It isspathaceous in shape, and of the same color andtexture as the corolla. This lineage contains onlyone other species, T. chrysea. The indumentum is oflepidote scales and small, glandular hairs (Fig. 1B).The wood is similar to that of Tabebuia group I butcontains tyloses, or intrusive growths of the cellwall into the vessel cavities, and pits intermediatein size between those of Tabebuia groups I and II(dos Santos and Miller 1992). The corolla is yellow(Fig. 1H), occasionally with thin red lines, andfruits are linear and irregularly costate and bullate.

TAXONOMIC CONSEQUENCES OF THE PHYLOGENY

The three clades of Tabebuia s. l. correspondclosely to the aggregations of species groups ofTabebuia discussed by Gentry (1992), and are

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FIG. 1. Characters defining clades of Tabebuia s. l. Photographs are arranged above the taxon they represent as indicated inbold on the tree. A–C. SEM’s of leaf or calyx indumentum. A. Handroanthus serratifolius, leaf undersurface showing long-stellate hairs and lepidote scales. B. Roseodendron donnell-smithii, calyx showing stalked glands. C. Tabebuia rosea, calyx showinglepidote scales. Scale bars for each photograph as indicated. D–F. Calyx shapes and flower color. D. Handroanthus chrysanthus

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recognized as genera here. The type of Tabebuia, T.cassinoides (Lamarck) A.P. de Candolle, was notincluded in the molecular study due to theunavailability of fresh material and the lack ofsuccess in extracting DNA from herbarium speci-mens. However, it has simple leaves and anindumentum of lepidote scales covering the leavesand calyx, which places it confidently in Tabebuiagroup I. We now restrict Tabebuia to those speciesoccurring in this clade. This corresponds toGentry’s (1992) informal groups 2 and 6–10 ofTabebuia. The sister taxon to this clade, Ekma-nianthe, shares the characters described for Tabebuiagroup I. It has 9-foliolate leaves, lepidote scales,and has similar wood anatomy; nevertheless, it ismorphologically distinct by virtue of its hawk-moth-pollinated flowers having long corolla tubeswith united lobes, and basally curved fruit. Theautapomorphy for Tabebuia I exclusive of Ekma-nianthe is its straight, cylindrical fruit.

The second clade, Tabebuia group II, comprisesmost of the species with yellow-flowers, andincludes many taxa segregated by Mattos (1970)into Handroanthus as well as Gentry’s (1992)informal groups 3–5. The type of Handroanthus,H. albus (Tabebuia alba), is not included in this studydue to lack of available tissue, but it shares the fourcharacters listed above and is assignable unambig-uously to this clade. Gentry (1972) insisted thatCouralia had nomenclatural priority over Han-droanthus as circumscribed by Mattos (1970).However, the type of Couralia, Tabebuia fluviatilis(Aubl) A.P. de Candolle, is assigned here toTabebuia I, so the name Couralia remains a synonymfor Tabebuia. Accordingly, priority rests withHandroanthus Mattos (Mattos 1970).

Tabebuia donnell-smithii was placed by Seibert(1940) in the genus Cybistax and later in Roseoden-dron by Miranda (1965). Roseodendron is resurrectedfor T. donnell-smithii and T. chrysea, since the latter,like T. donnell-smithii has calyces that are texturallyidentical to the corolla. These two species formGentry’s (1992) informal group 1.

TAXONOMIC DESCRIPTIONS

HANDROANTHUS Mattos, Loefgrenia 50: 2 1970.—TYPE: Handroanthus albus (Cham.) Mattos.

Trees or occasionally shrubs; heartwood distinct

from sapwood, olive brown to blackish, lapacholpresent in large quantities; Rays 1–3 cells wide,intervessel pits large (8–14 mm), fibers thick walled;wood, very dense with high specific gravity (,7.4). Leaves (3)5–9 foliolate (reported to be occa-sionally 1-foliolate in H. pumilus and H. selachiden-tatus), leaflets narrowly elliptic to broadly ovate(17 cm long and 18.5 cm wide), with simple,forked, stellate, barbate or dendroid trichomes atleast along vein axils and sometimes denselycovering leaves; petiolule to 9 cm; petiole to30 cm. Inflorescence dichotomously branched,without a well developed central rachis, sometimescontracted; pubescence with simple, stellate, bar-bate or dendroid trichomes. Flowers: Calyx co-riaceous, campanulate 5-dentate, 4–20 mm long, 3–20 mm wide; trichomes simple, stellate or den-droid, sometimes forming a dense covering. Co-rolla yellow or magenta with yellow throat,tubular-infundibuliform to tubular-campanulate,tube 2.5–6.5 cm long, 0.6–3.5 cm wide at mouth,lobes 0.5–5 cm; tube glabrous to densely tomentosewith stellate to dendroid or barbate trichomes;lobes sometimes pilose in sinuses; throat papillose,pubescent at level of stamen insertion. Stamens

didynamous; thecae divaricate, 1.5–2 mm (20 mmin H. subtilis) staminode reduced. Ovary conical tolinear-oblong; ovules 2–10 seriate in each locule.Fruit an elongate linear to cylindric capsule,smooth to slightly costate, glabrous to scatteredlepidote to pubescent; pubescence scattered tovillous; trichomes simple, stellate or dendroid.Seeds thin, bialate; wings hyaline membranacousand sharply demarcated from seed body.

A genus of 30 species distributed throughoutCentral and South America with one species (H.billbergii) in the Antilles.

Taxa transferred to Handroanthus (*indicates thenine taxa used in Grose and Olmstead 2007):

1. HANDROANTHUS ALBUS (Chamisso) Mattos, Loef-grenia 50: 2. 1970. Tecoma alba Chamisso,Linnaea 7: 655. 1832. Tabebuia alba (Chamisso)Sandwith, Lilloa 14: 136. 1948.—TYPE: BRA-ZIL. Parana and Rio Grande do Sul, Sellow s.n.(B, lectotype HBG).

2. H. arianeae (A. H. Gentry) S. Grose, comb. nov.Tabebuia arianae Gentry, Flora Neotropica25(2): 142. 1992.

r

ssp. chrysanthus- cupular 5-dentate calyx covered with red pubescence, and yellow corolla E. Roseodendron donnell-smithii,spathaceous calyx yellow, and the same texture and color as the corolla. F. Tabebuia platyantha, bilabiate calyx, white corolla.Photo by A. H. Gentry courtesy of Tropicos3. G–I. Flower color (see also D–F). G. Handroanthus impetiginosus, pink corollas withyellow throat. H. Roseodendron donnell-smithii, yellow corolla. I. Tabebuia haemantha, fuchsia corolla. All photos, by S.O. Grose,unless otherwise indicated. Phylogeny from (Grose and Olmstead 2007).

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3. H. BARBATUS (E. Mey.) Mattos, Loefgrenia 50: 2.1970. Bignonia barbata E. Mey. Nova ActaPhys.-Med. Acad. Caes. Leop.-Carol. Nat.Cur. 12: 782. 1825. Zeyheria barbata (E. Meyer)Miquel. Flora 25: 430. 1842. Tecoma barbata (E.Meyer) DC. Prodr. 9: 221. 1845. Tabebuiabarbata (E. Meyer) Sandwith, Lilloa 3: 462.1938.

Bignonia fluviatilis Aublet sensu Humboldt, Bon-pland and Kunth, Nov. gen. sp. Pl3: 139. 1819non Aublet.

Tecoma toxophora Martius emend DC, Prodr. 9: 217.1845 (excl. sin. Margr.) non Martius, Flora 24,Beibl. 15. 1841. Couralia toxophora (Martiusemend DC) Bentham and Hooker f. ex K.Schum., Nat. Pflanzenfam. 4(3b): 239. 1894.

4. H. billbergii (Bur. & K.Schum.) S. Grose, comb.nov. Tecoma billbergii Buerau & K. Schum. inMartius, Fl. Bras. 8(2): 319. 1897.

a. H. billbergii. subsp. billbergii. Tabebuia bill-bergii subsp. billbergii Bureau & K. Schum. inMartius, Fl. Bras 8(2): 319. 1897.

b. H. billbergii subsp. ampla (A. H. Gentry) S.Grose, comb. nov. Tabebuia billbergii subsp.ampla A. Gentry, Phytologia 35: 187. 1977.

Tabebuia ecuadoriensis Standley, Trop. Woods 46:17. 1936.

5. H. botelhensis (A. H. Gentry) S. Grose, comb.nov. Tabebuia botelhensis A. Gentry, FloraNeotropica 25(2): 150. 1992.

6. H. bureavii (Sandwith) S. Grose, comb. nov.Tabebuia bureavii Sandwith, Kew Bull. 1958:442. 1992. Tecoma dentata Bur. & K. Schum,Martius Fl. Bras. 8(2): 323. 1897, not Tabebuiadentata, 1863. Handroanthus dentatus (Bur. andK. Schum.) Mattos, Loefgrenia 50: 2. 1970.

7. H. CAPITATUS (Bureau ex. K. Schum.) Mattos,Loefgrenia 50: 2. 1970. Tabebuia capitata (Bu-reau ex K. Schum.) Sandwith, Rec. Trav. Bot.Neerl. 34: 226. 1937. Tecoma capitata Bureau &K. Schum., Fl. Bras. 8(2): 337. 1897.

Tecoma leucoxylon (L.) Mart. ex. DC var. miqueliiDC, Prodr. 9: 219. 1845.

Tabebuia glomerata Urban, Feddes Rebert. 14: 305.1916.

Tabebuia hypolepra Sprague and Sandwith, KewBull. 1932: 25 1932.

8. H. catarinensis (Gentry) S. Grose comb. nov.Tabebuia catarinensis Gentry, Ann. MissouriBot. Gard. 64: 318. 1977.

9. H. chrysanthus (Jacq.) S. Grose comb. nov.Bignonia chrysantha Jacq., Pl. hort. Schoenbr.2: 45, tab. 211. 1797. Tecoma chrysantha (Jacq)DC, Prodr. 9: 211. 1845. Tabebuia chrysantha(Jacq.) Nichols. subsp. Chrysantha, Ill. dict.gard. 4: 1. 1887.

a. H. chrysanthus subsp. chrysanthus*

Tabebuia rufescens J. R. Johnston, Proc. Amer.Acad. Arts 40: 696. 1905.

Tecoma evenia Donn. Sm., Bot. Gaz. (Crawfordsville) 20. 1895 pro parte.

Tecoma palmeri Kraenzl. Feddes Repert, 17: 220.1921.

b. H. chrysanthus subsp. meridionalis (A. H.Gentry) S. Grose, comb. nov. Tabebuiachrysantha ssp. meridionalis A. Gentry, Phy-tologia 35: 193. 1977.

Tecoma spectabilis Planchon and Linden Fl. SerresJard. L’Europe. 9: 233. 1854.

Tabebuia spectabilis (Planch. and Linden) Nichols., Ill. dict. gard. 4: 1. 1887.

c. H. chrysanthus subsp. pluvicolus (A. H.Gentry) S. Grose comb. nov.* Tabebuiachrysantha (Jacquin) Nicholson subsp. pluvi-cola. A. Gentry in Phytologia 35: 190. 1977.

Tecoma grandis Appun., Behand. Samereien undPflanzen 39. 1858, nom. nud.

10. H. CHRYSOTRICHUS (Mart. ex DC) Mattos,Loefgrenia 50: 2. 1970.* Tecoma chrysotrichaMart. ex. DC, Prodr. 9: 216. 1845. Gelseminumchrysotrichum (Mart. ex. DC) O. Kuntze, Rev.gen. pl. 3(2): 245. 1898.

Tecoma ochracea var. denudata Chamisso, Linnaea 7:653. 1832.

Tecoma flavescens Mart. ex. DC, Prodr. 9: 216. 1845.

Tecoma obstusata DC, Prodr. 9: 217. 1845.Tecoma chrysotricha var. obtusata (DC) Bur. & K.

Schum, Martius Fl. Bras. 8(2): 338. 1897.Tabebuia chrysotricha (Mart. ex. DC Standley,Publ. Field. Mus. Nat. Hist. Bot. Ser. 11: 176.1936. Tabebuia chrysothrica var. obtusata(DC)Toledo, Arq. bot. estado Sao Paulo 3(1):35. 1952. Handroanthus chrysotrichus var. obtu-sata (DC) Mattos, Loefgrenia 50: 2. 1970.

Tecoma pedicellata Bur & K. Schum., Martius Fl.Bras. 8(2): 336. 1897. Handroanthus pedicellatus(Bureau & K. Schum.) Mattos, Loefgrenia 50: 2.1970.

Tecoma grandis Kraenzl., Feddes Repert. 17: 217.1921

11. H. coralibe (Standley) S. Grose comb. nov.Tabebuia coralibe Standley in Trop. Woods. 36:18. 1933.

12. H. cristatus (Gentry) S. Grose comb. nov.Tabebuia cristata A. Gentry in Flora Neotropica25(2): 174. 1992.

13. H. guayacan (Seeman) S. Grose comb. nov.*Tecoma guayacan Seemann, Bot. voy. Herald180. 1854. Tabebuia guayacan (Seemann) Hems-ley in Biol. Centr.-Amer Bot. 2: 495. 1882.

14. H. HEPTAPHYLLUS (Vell.) Mattos, Loefgrenia 50:2. 1970. Bignonia heptaphylla Vell., Fl. Flumin.

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251. 1829. Tabebuia heptaphylla (Vell.) Toledo,Arq. Bot. Estado Sao Paulo, n. s. 3: 33. 1952.

Tecoma eximia Miq., Linnaea 22: 803. 1849. Tabebuiaeximia (Miq.) Sandwith, Lloydia 2: 213. 1939.Handroanthus eximia (Miq.) Mattos, Loefgrenia50: 2. 1970.

Tecoma curialis Saldanha da Gama, Config, descr.orgaos fund. de Janeiro. 1: 51. 1865.

Tecoma ipe Mart. ex. K. Schum., Engl. and Prantl,Pflanzenfam. 4(3b): 238. 1894. Tabebuia ipe(Mart. ex. K. Schum.) Standley, Trop. Woods.36: 20. 1933.

Tecoma ipe var. desinens Sprague, Bull. Herb.Boissier, ser. 2, 5: 86. 1905.

Tecoma ipe var. desinens f. parviflora Sprague, Bull.Herb. Boissier, ser. 2, 5: 86. 1905.

Tecoma ipe var. desinens f. grandiflora Sprague, Bull.Herb. Boissier, ser. 2, 5: 86. 1905.

Tabebuia avellanedae var. paulensis Toledo, Arq. Bot.Estado Sao Paulo, n. s., 3: 1952.

Tabebuia impetiginosa var. lepidota (Bur.) Toledo,Arq. Bot. Estado Sao Paulo, n. s., 3: 1952.Handroanthus impetiginosus var. lepidotus (Bur.)Mattos, Loefgrenia 50: 2. 1970.

15. H. IMPETIGINOSUS (Mart. Ex DC) Mattos, Loef-grenia 50: 2. 1970.* Tecoma impetiginosa Mart.ex. DC, Prodr. 9: 218. 1845. Tabebuia impetigi-nosa (Mart. ex. DC) Standley, Publ. Field Mus.Nat. Hist., Bot. Ser. 11: 176. 1936.

Tabebuia avellanedae Lorentz ex. Griseb., Symbol. fl.argent. 258. 1879. Gelseminum avellanedae (Lor-entz ex Grisebach) Kuntze, Rev. gen. pl. 3(2):245. 1898. Handroanthus avellanedae (Lorentz exGriseback) Mattos, Loefgrenia 50: 2. 1970.

Tabebuia palmeri Rose, Contr. U. S. Natl. Herb. 1:109. 1891.

Tecoma impetiginosa var lepidota Bureau, Vidensk.Meddel. Dansk. Naturhist. Foren. Kjøbenhavn.1893: 114. 1894. Handroanthus impetiginosusvar. lepidotus (Bureau) Mattos, Loefgrenia 50:2. 1970.

Tecoma adenophylla K. Schum. ex. Bureau & K.Schum. in Martius, Fl. Bras. 8(2): 412. 1897.

Tecoma ipe var. integra Sprague, Bull. Herb. Biossier,ser. 2 5: 86. 1905. Tecoma integrum (Sprague)Chodat, Bull. Soc. Bot. Geneve ser. 2, 9: 242.1917. Tabebuia ipe var. integra (Sprague) Sand-with, Lloydia 2: 213. 1939.

Tecoma ipe var. integrifolia Hassler, Rev. Inst. Parag.3: 166. 1901.

Tecoma avellanedae var. alba Lillo, Seg. contr. arb.Argent. 13. 1917.

Tabebuia nicaraguensis Blake, Contr. Gray Herb. 52:95. 1917.

Tabebuia dugandii Standley, Trop. Woods 36: 17.1933.

Tabebuia schunkevigoi Simpson, Fieldiana, Bot. 36: 1.1972.

16. H. incanus (A. H. Gentry) S. Grose comb. nov.Tabebuia incana A. Gentry in Ann. MissouriBot. Gard. 65: 732. 1978.

17. H. lapacho (K. Schum.) S. Grose comb. nov.Tecoma lapacho K. Schum., Pflanzenfam. 4(3b):238. 1894. Tabebuia lapacho (K. Schum.) Sand-with, Lilloa 14: 136. 1948.

Tabebuia flavescens (Vell.) Griseb., sensu Griseb.,Symb. fl. Argent. 257. 1879.

18. H. OBSCURUS (Bur. ex. K Schum.) Mattos,Loefgrenia 50: 2. 1970.* Tecoma obscura Bur.ex K. Schum., Mart, Fl. Bras. 8(2): 343. 1897.Tabebuia obscura (Bureau ex. K. Schum.) Sand-with, Rec. Trav. Bot. Neerl. 34: 226. 1937.

Tabebuia subtilis var. schultesiana Sandwith, Bot.Mus. Leafl. Harvard Univ. 17: 96. 1955.Tabebuia obscura var schultesiana (Sandwith)Sandwith in Dugand, Mutisia 25: 16. 1956.

19. H. OCHRACEUS (Cham.) Mattos, Loefgrenia 50:2. 1970.

a. H. OCHRACEUS subsp. HETEROTRICHUS (DC) S.Grose. Tecoma heterotricha DC, Prodr. 9: 219.1845. Tabebuia heterotricha (DC) Hemsley,Biol. Centr.-Amer., Bot. 2: 495. 1882. Tabe-buia ochracea (Cham) Standley subsp. hetero-trichus (DC) A. H. Gentry, Fl. Venezuela8(4): 391. 1982.

b. H. OCHRACEUS subsp. NEOCHRYSANTHUS (A. H.Gentry) S. Grose.* Tabebuia neochrysantha A.H. Gentry, Brittonia 22: 260. 1970. Tabebuiaochracea (Chamisso) Standley subsp. neo-chrysantha (A. H. Gentry) A. H. Gentry,Ann. Missouri Bot. Gard. 60: 948. 1974.

Tabebuia chrysantha (Jacq.) Nichols., Dict. Gard. 4: 1.1897, sensu Sandwith, non Jacq.

c. H. OCHRACEUS (Cham.) Mattos subsp. OCHRA-

CEUS. Tecoma ochracea Cham., Linnaea 7: 653.1832.

Tecoma hypodictyon DC, Prodr. 9: 217. 1845. Tabebuiahypodiction (DC) Standl., Field Mus. Nat. Hist.,Bot. Ser. 11: 176. 1936.

Tecoma heteropoda DC, Prodr. 9: 219. 1845. Tabebuiaochracea subsp. heteropoda (DC) A. H. Gentry,in Prance, Biological Diversification in theTropics 132. 1982.

Bignonia tomentosa Pav. ex DC, Prodr. 9: 219. 1845.nom. nud., pro syn.

Tecoma hassleri Sprague, Trans. Proc. Bot. Soc.Edinburgh 48: 435. 1904.

Tecoma grandiceps Kranzl., Feddes Repert. 17: 216. 1921.

Tecoma campinae Kraenzlin, Feddes Repert. 17: 215.1921.

Tecoma hemmendorffiana Kraenzl., Feddes Repert.17: 224. 1921.

2007] GROSE & OLMSTEAD: REVISION OF TABEBUIA 665

20. H. PEDICELLATUS (Bur. ex K. Schum.) Mattos,Loefgrenia 50: 2. 1970. Tecoma pedicellata Bur. &K. Schum., in Mart., Fl. Bras. 8(2): 336. 1897.Tabebuia pedicellata (Bur. ex. K. Schum.) A. H.Gentry, Flora Neotropica 25(2): 236. 1992.

Tecoma catinga Bur. & K. Schum., in Martius, Fl.Bras. 8(2): 337. 1897. Handroanthus catinga (Bur.& K. Schum.) Mattos. Loefgrenia 50: 4. 1970.

21. H. pulcherrimus (Sandwith) S. Grose comb.nov. Tabebuia pulcherrima Sandwith, Lilloa 14:133. 1948. Tecoma petropolitana Glaz. nom.nud., Mem. Soc. Bot. France 3: 528. 1911.

22. H. pumilus (A. H. Gentry) S. Grose comb. nov.Tabebuia pumila A. H. Gentry, Flora Neotropica25(2): 244. 1992.

23. H. riodocensis (A. H. Gentry) S. Grose comb.nov. Tabebuia riodocensis A. H. Gentry, FloraNeotropica 25(2): 1248. 1992.

24. H. selachidentatus (A. H. Gentry) S. Grosecomb. nov. Tabebuia selachidentata A. H. Gen-try, Flora Neotropica 25(2): 254. 1992.

25. H. serratifolius (Vahl) S. Grose comb. nov.*Bignonia serratifolia Vahl, Eclog. Amer. 2: 46.1798. Tecoma serratifolia (Vahl) G. Don., Gen.syst. 4: 224. 1838. Tabebuia serratifolia (Vahl)Nichols. in Dict. Gard. 4: 1. 1887.

Bignonia flavescens Vell., Fl. flumin. 252. 1829 (1825).Tecoma flavescens (Vell.) Martius ex DC, Prodr.9: 226. 1845. Handroanthus flavescens (Vell.)Mattos, Loefgrenia 50: 2. 1970.

Bignonia araliacea Cham., Linnaea 7: 683. 1832.Tecoma araliacea (Cham.) DC, Prodr. 9: 221.1845. Tabebuia araliacea (Cham.) Morong andBritton, Ann. N. Y. Acad. Sci. 7: 190. 1893.Gelseminum araliaceum (Cham.) O. Kuntze,Rev. gen. 3(2): 245. 1898. Handroanthus aralia-ceus (Cham.) Mattos, Loefgrenia 50: 2. 1970.

Tecoma conspicua DC, Prodr. 9: 221. 1845. Bignoniaconspicua Richard ex DC, Prodr. 9: 221, nom.nud., pro syn.

Tecoma patrisiana DC, Prodr. 9: 221. 1845.

Tecoma speciosa DC, Prodr. 9: 218. 1845. Gelseminumspeciosum (DC) O. Kuntze, Rev. gen. 3(2): 245.1898.

Tecoma nigrescens Klotz in Schomburgk, Reisen 3:1159. 1848, nom. nud.

Tecoma atractocarpa Bur. & K. Schum., Martius, Fl.Bras. 8(2): 326. 1897.

Handroanthus atractocarpus (Bur. & K. Schum.)Mattos, Loefgrenia 50: 2. 1970.

Tabebuia monticola Pittier, Cat. Flora Venez. 2: 409.1947, nom. nud.

26. H. spongiosus (Rizzini) S. Grose comb. nov.Tabebuia spongiosus Rizz., Rodriguesia 28: 172.1976.

27. H. subtilis (Sprague & Sandwith) S. Grose

comb. nov.* Tabebuia subtilis Sprague andSandwith, Kew Bull. 1932: 23. 1932.

28. H. uleanus (Kraenzl) S. Grose comb. nov.Tecoma uleana Kranzlin, Feddes Repert. 17:217. 1921. Tabebuia uleana (Kraenzl.) A. H.Gentry in Mem. N.Y. Bot. Gard. 29: 279. 1978.

29. H. UMBELLATUS (Sonder) Mattos, Loefgrenia 50:2. 1970.* Tecoma umbellata Sond., Linnaea 22:562. 1849. Tabebuia umbellata (Sond.) Sandwith,Lilloa 14: 136. 1948.

Tecoma eximia Miq., Linnaea 22: 803. 1849. Tabebuiaeximia (Miq.) Sandwith, Lloydia 2: 213. 1939.Handroanthus eximius (Miq.) Mattos, Loefgre-nia 50: 2. 1970.

Tecoma umbellata var. lanceolata Bur. & K. Schum.,Martius, Fl. Bras. 8(2): 335, 1897. Tabebuia um-bellata var. lanceolata (Bur. & K. Schum.) Toledo,Arq. Bot. Estado Sao Paulo 3(1): 35. 1952.Handroanthus umbellatus var. lanceolatus (Bur. &K. Schum.) Mattos, Loefgrenia 50: 2. 1970.

30. H. VELLOSOI (Toledo) Mattos, Loefgrenia 50: 2.1970. Tabebuia vellosoi Toledo, Arq. Bot. EstadoSao Paulo, n. s. 3(1): 34. 1952.

Bignonia longiflora Vell., Fl. flumin. 252. 1829 (1825).Tecoma longiflora (Vell.) Bur. & K. Schum., inMartius, Fl. Bras. 8(2): 324. 1897, non Tecomalongiflora Griseb. 1866.

Tecoma alba Cham. var. subdenudata Bur., Dansk.Naturhist. Foren. Kjøbenhavn 1893: 115. 1893.

ROSEODENDRON Miranda, Bol. Soc. Not. Mex. 29:431965.—TYPE: Roseodendron donnell-smithii(Rose) Miranda.

Trees to 35 m; heartwood not very distinct fromsapwood, yellowish or light to reddish brown;tyloses present; lapachol present in small quanti-ties (R. chrysea); rays 2–5(8) cells wide; mediumintervessel pits (6–8 mm); thin to thick walledfibers; medium specific gravity (0.4–7.4). Leaves

palmately 5–7 foliolate; leaflets oblong-elliptic, to28 cm long and 14 cm wide, usually much smaller,membranaceous to chartaceous, the surface some-what rough, especially above, sometimes puber-ulous above and below with simple or dendroidtrichomes, or more or less glabrate, drying olive,darker above and lighter below; apex acute toacuminate; bases truncate; petiolules to 7 cm long;petiole to 26 cm long, tomentose with dendroidand barbate trichomes or glabrate. Inflorescence

a large terminal panicle with the central rachis welldeveloped and lateral branches either very shortand dendroid-pubescent, or branching with rachisand branches puberulous with capitate trichomes.Flowers: Calyx campanulate, 10–20 mm long, 5–15 mm wide, finely membranaceous, bilabiate toirregularly several–5-dentate, somewhat lepidote

666 SYSTEMATIC BOTANY [Volume 32

and puberulous, sometimes with gland tippedtrichomes. Corolla yellow, sometimes with reddishlines in throat, tubular-infundibuliform; tube 3–4.5 cm long, 0.8–2 cm wide at mouth of tube, lightlypuberulous to almost glabrous outside, generallyglabrous inside, pubescent at level of stameninsertion; lobes glabrous to lightly pubescent; lobes1–2.5 cm long. Stamens didynamous; thecae di-varicate, 2–2.5 mm long; staminode reduced. Ovarylinear, 5–6 mm long, 0.6–1.5 mm wide; ovules 4 or 6seriate in each locule. Fruit an elongate-linearcapsule, 25–45 cm long, 0.9–3 cm wide, longitudi-nally striate-costate to irregularly striate costate with8–12 rather irregular longitudinal ribs, puberulouswith simple and branched or simple-gland-tippedtrichomes. Seeds thin; wings hyaline-membranac-eous and sharply demarcated from seed body.

Two species ranging from Mexico to Northwest-ern South America, and dry areas of Venezuelaand Colombia.

Taxa transferred to Roseodendron (* indicatestaxon used in Grose and Olmstead, 2007):

1. ROSEODENDRON CHRYSEUM (Blake) Miranda, Bol.Soc. Mex. 29: 43. 1965. Tabebuia chrysea Blake,Contr. Gray herb. 53: 50. 1918. Tecoma chrysea(Blake) Pittier, publ. Pl. Usual. Venez. 63. 1939.Cybistax chrysea (Blake) Seibert, Trop Woods63: 7. 1940.

2. R. DONNELL-SMITHII (Rose) Miranda, Bol. Soc.Mex. 29: 43. 1965.* Tabebuia donnell-smithiiRose, Bot. Gaz. 17: 418. pl. 26. 1892. Cybistaxdonnell-smithii (Rose) Seibert, Carnegie Inst.Wash. Publ. 522: 392. 1940.—TYPE: GUATE-MALA. Escuintla, Donnell-Smith 2070 (US!).

Cybistax millsii Miranda, Bol. Soc. Mex. 26: 129.1961, Roseodendron millsii (Miranda) Miranda,Bol. Soc. Mex. 29: 43. 1965, Tabebuia millsii(Miranda) A. H. Gentry, Ann. Missouri Bot.Gard. 63: 75. 1976.

TABEBUIA Gomes ex A.P. de Candolle, Biblioth.Universelle Geneve, ser 2, 17:130. Sep. 1838.—TYPE: Tabebuia uliginosa (Gomes) DC 5

Tabebuia cassinoides (Lam.) DC, Encyc. Meth.1: 418. 1785.

Leucoxylon Raf., Sylva Tellur. 77. Oct 1838.—TYPE:L. riparia Raf. 5 T. heterophylla (DC) Britton.

Potamoxylon Raf., Sylva Tellur. 78. Oct 1838.—TYPE: P. alba Raf. 5 T. fluviatilis (Aubl. ) DC.

Proterpia Raf., Sylva Tellur. 80. Oct 1838.—TYPE: P.obtusifolia (Lamarck) Raf. (Bignonia obtusifoliaLam.) 5 T. cassinoides (Lamarck) DC

Couralia Splitgerberger, Tujdschr. Natuurl. Gesch.Physiol. 9: 14. 1842.—TYPE: Couralia fluviatilis(Aubl.) Splitgerberger 5 T. fluviatilis (Aubl.) DC.

Trees or shrubs; heartwood not distinct fromsapwood, light to reddish brown; lapachol absent;rays 1–3 cells wide; small to medium intervesselpits (3–6 mm); thin to thick walled fibers; mediumspecific gravity (0.4–7.4). Leaves 1–7(9) foliolate;leaflets narrowly elliptic to orbicular to 35 cm longand 32 cm wide, with stalked or sessile lepidotescales that are usually scattered and sometimesdensely covering undersurface of leaves; petioluleto 11 cm; petiole to 18 cm. Inflorescence dichoto-mously branching, without a well developed centralrachis, usually a few flowered panicle, occasionallymany flowered, sometimes densely lepidote. Flow-ers: Calyx coriaceous, 2–3 labiate rarely 5-dentate,5–25 mm long, 4–11 mm wide, densely lepidote.Corolla white to red, often with yellow throat,completely yellow in two species (T. nodosa and T.aurea) tubular-infundibuliform to tubular-campan-ulate; tube 2–7 cm long, 0.6–3.5 cm wide at mouth;lobes 0.5–3.2; tube glabrous to pubescent at level ofstamen insertion. Stamens didynamous; thecaedivaricate, 2–6 mm; staminode reduced. Ovarylinear; ovules 2–3 seriate in each locule. Fruit anelongate linear to cylindric capsule, smooth tocostate striate, minutely to densely lepidote. Seedsthin, bialate; wings hyaline membranacous andsharply demarcated from seed body.

A genus of 67 species widely distributedthroughout Central and South America and theAntilles.

Included taxa (* indicates taxon used in Groseand Olmstead 2007). Names in this group remainunchanged from Gentry (1992) and are listed forconvenience:

1. TABEBUIA ACROPHYLLA (Urb.) Britton*2. T. ANGUSTATA Britton3. T. ARIMAOENSIS Britton4. T. AUREA (Silva Manso) Benth. & Hook. f. ex S.

Moore*5. T. BAHAMENSIS (Northr.) Britton*6. T. BERTEROI (DC) Britton*7. T. BIBRACTEOLATA (Griseb.) Britton8. T. BROOKSIANA Britton9. T. BUCHII (Urb.) Britton10. T. BULLATA A. H. Gentry11. T. CALCICOLA Britton12. T. CALETICANA A. H. Gentry and D. Albert13. T. CASSINOIDES (Lam.) DC (Type)14. T. CLEMATIS Alain15. T. CONFERTA Urb.16. T. CRISPIFLORA Alain17. T. XDEL-RISCOI Borhidi18. T. DENSIFOLIA Urb.19. T. DOMINGUENSIS (Urb.) Britton20. T. DUBIA (C.Wright ex Sauvalle) Britton ex

Seibert

2007] GROSE & OLMSTEAD: REVISION OF TABEBUIA 667

21. T. ELEGANS (Urb.)22. T. ELLIPTICA (DC) Sandwith23. T. FLUVIATILIS (Aubl.) DC24. T. GLAUCESCENS Urb.25. T. HAEMANTHA (Bertero ex Sprengel) DC*26. T. HETEROPHYLLA (DC) Britton*27. T. HYPOLEUCA (Wright ex Sauvalle) Urb.28. T. INAEQUIPES Urb.29. T. INSIGNIS (Miq.) Sandwith

a. T. INSIGNIS (Miq.) Sandwith var. INSIGNIS*b. T. INSIGNIS (Miq.) Sandwith var. MONO-

PHYLLA Sandwithc. T. INSIGNIS (Miq.) Sandwith var. PACIMO-

NENSIS Sandwith

30. T. JACKIANA Ekman ex Urb.31. T. LEPIDOPHYLLA (A. Richard) Greenm.32. T. LEPIDOTA (HBK) Britton*33. T. LEPTONEURA Urb.34. T. LINEARIS Alain35. T. MAXONII Urb.36. T. MICROPHYLLA (Lam.) Urb.*37. T. MOAENSIS Britton38. T. MULTINERVIS Urb. and Ekman39. T. MYRTIFOLIA (Griseb.) Britton

a. T. MYRTIFOLIA (Griseb.) Britton var. MYR-

TIFOLIA

b. T. MYRTIFOLIA (Griseb.) Britton var. PET-

ROPHILA (Greenm) A. H. Gentry

40. T. NODOSA (Griseb.) Griseb.41. T. OBOVATA Urb.42. T. OBTUSIFOLIA (Cham.) Bur.43. T. OPHIOLITICA Alain44. T. ORINOCENSIS (Sandwith) A. H. Gentry45. T. PALLIDA (Lindl.) Miers46. T. PALUSTRIS Hemsl.*47. T. PANICULATA Leonard48. T. PILOSA A. H. Gentry49. T. PINETORUM Britton

50. T. PLATYANTHA (Griseb.) Britton

51. T. POLYANTHA Urb. & Ekman

52. T. POLYMORPHA Urb.

53. T. PULVERULENTA Urban

54. T. RETICULATA A. H. Gentry

55. T. REVOLUTA (Urb.) Britton

56. T. RIGIDA Urb.

57. T. ROSEA (Bertol.) DC*

58. T. ROSEO-ALBA (Ridl.) Sandwith

59. T. SAUVALLEI Britton*

60. T. SCHUMANNIANA Urb.*

61. T. SHAFERI Britton

62. T. SIMPLICIFOLIA Carabia ex Alain

63. T. STENOCALYX Sprague & Stapf

64. T. STRIATA A. H. Gentry*

65. T. TRACHYCARPA (Griseb.) K.Schum.

66. T. VINOSA A. H. Gentry

67. T. ZANONII A. H. Gentry

Tabebuia Alliance. The character uniting themembers of the Tabebuia alliance and distinguish-ing them from other Bignoniaceae is digitately-compound leaves. Members of this alliance areusually shrubs to large trees, and the leaves areoften covered with lepidote scales and sometimeswith simple or branched hairs. The flowers havea calyx that is spathaceous or cupular with 3–5lobes. The calyx and corolla often have anindumentum of lepidote scales or hairs. The ovaryis linear or ovate, and bilocular at least in extremebase. The fruit has striations, ridges or spines, andis usually loculicidally dehiscent but is indehiscentin the tree genera of the Crescentieae. The seeds ofwind-dispersed taxa are winged, those that arewater or mammal dispersed have vestigial wings.

Key to the Genera. Not all specimens arecollected at a time of year to show all of thedesired features, usually a combination of leaves,flowers, and fruit. This key is intended to be alsouseful for specimens that are sterile or contain onlyflowers and fruits. The most distinguishing fea-tures are in bold.

1a. Young branches and inflorescence appear shiny with sticky secretions. Leaves thin, membranaceous in texture,glandular, drying dark. Calyx similar in texture to corolla, but different in color; corolla white to pale pink, sometimeswith darker lines in throat. Fruit linear, terete capsule, valves subwoody. Seeds linear, seed wings made of minutehairs. One sp. Continental South America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sparattosperma

1b. Young branches and inflorescence not shiny with sticky secretions, instead with indumentum of hairs and/or lepidotescales. Leaves various, sometimes with plate shaped glands. Calyx, corolla, and fruits various. Widespread . . . . . 2

2a. Leaves, young branches and inflorescence with indumentum of simple, branched or stellate hairs, never stalkedlepidote, sessile lepidote scales usually present, plate shaped glands sometimes present. Leaves 3–9 foliolate.Flowers yellow or brown, rarely magenta. Widespread . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3a. Leaves 5–9 foliolate, glandular punctate, with plate-shaped glands on both surfaces; usually with simple hairs(however occasionally glabrous). Inflorescence a terminal raceme, covered with simple hairs, flowers small,yellow/brown occasionally with mauve in throat. Fruit linear, spirally coiled. Seeds bialate with hyalinewings. 2 spp. Dry forests in Central and South America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Godmania

3b. Leaves 3–9 foliolate, not glandular punctate, with indumentum of stellate or branched hairs, often denselycovering surface. These hairs also present on new growth. Inflorescence various often with stellate hairs,especially on calyx and ovary. Flowers and fruit various . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

668 SYSTEMATIC BOTANY [Volume 32

4a. Plants shrubs or subshrubs, to 3 m tall. Leaves 5-foliolate, with indumentum of dense stellate or dendroidtomentose hairs, especially on lower surface of leaf, calyx and ovary, and sometimes corolla. Corolla,yellow/brown, sometimes pinkish in throat; calyx bilabiate, split to base, densely pubescent withstellate hairs. Fruit flattened orbicular, with spiny projections, also densely stellate tomentose. Seedsorbicular. 2 spp. Restricted to Brazilian Cerrado . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Zeyheria

4b. Plants usually trees, rarely shrubs. Leaves 3–9 foliolate, reportedly 1-foliolate in one Brazilian species, withindumentum of simple, dendroid or stellate hairs. These hairs often tannish, sometimes reddish or white.Flowers usually yellow often with red lines in throat, rarely magenta, tubular campanulate, calyxusually 5 dentate, rarely spathaceous, covered with hairs, sometimes densely so. Fruit a linear capsule;seeds linear with hyaline wings. 30 spp. Continental Central and South America. . . . . . . Handroanthus

2b. Leaves, young branches and inflorescence with indumentum of stalked or sessile lepidote scales, if hairs present,simple (decapitated stalked lepidote?). Leaves simple to 9 foliolate. Flowers rarely yellow, usually greenish white orwhite to red. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

5a. Inflorescences cauliflorous or ramiflorous, growing from round or branchlike areas on bark (terminal in somespecies of Amphitecna) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

6a. Fruit dehiscent, linear, sometimes curved or coiling; septum thickened with pits in which seeds rest. Seeds verysmall. Leaves simple or 3–7 foliolate, shiny when dry, drying dark, undersurface with simple hairs formingdomatia, these drying contrastingly lighter than leaf, if domatia absent, then leaves simple. Leaf bases truncate,apices cordate to apiculate. Petioles usually at least 2 cm, petiolules at least 1 cm. Inflorescences ramiflorous orcauliflorous; calyx tubular-campanulate, irregularly dentate; flowers with pedicel elongate; corolla yellow todark red, lower corolla lobes reflexed; stamens exserted, 4 spp. Cuba and Hispanola . . . . . . . . . . Spirotecoma

6b. Fruit indehiscent, linear to spherical, seeds wingless, or with vestigial wings. Leaves usually simple, whencompound 3–5 foliolate, not drying as above, membranaceous, glabrous or lepidote, sometimes with stipularspines. Inflorescences cauliflorous, ramiflorous or terminal; flowers campanulate with large bulge in throat,corolla greenish white sometimes with red or purple lines. (Crescentieae) . . . . . . . . . . . . . . . . . . . . . . . . . 7

7a. Leaves 3–5 foliolate, often with stipular spines. Fruit a linear capsule, bilocular throughout, seeds smallwith vestigial wings, no pulp in fruit. Flowers terminal and cauliflorous, greenish white; calyxmembranaceous, irregularly spathaceously split. 8 spp. Mexico to extreme NW Colombia . . Parmentiera

7b. Leaves simple (in part trifoliolate in Crescentia alata). Fruit a pepo, bilocular to approximately halfway.Seeds embedded in white fleshy, sweet pulp. Inflorescences cauliflorous or terminal, calyx thick, tearingirregularly. Central America and Greater Antilles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

8a. Leaves simple, often with a red pulvinus on short petiole. Inflorescence terminal or cauli/ramiflorous.Flowers greenish or creamy white, petal lobes united into rim. Fruits oblong (if orbicular, thenmangrove species). Seeds very large, 2 cm wingless. 22 spp. wet forests, Central America Amphitecna

8b. Leaves usually simple, trifoliolate with winged rachis in C. alata, linear in C. linearifolia pulvinus woody,if present. Inflorescences cauliflorous. Flowers greenish white, sometimes with red lines; petal lobeswith long, drooping apices. Fruit, green, orbicular. Seeds very small and numerous. 6 spp. CentralAmerica and Greater Antilles, one species cultivated worldwide (C. cujete) . . . . . . . . . . . Crescentia

5b. Inflorescences terminal or axillary never cauliflorous or ramiflorous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99a. Calyx campanulate, 5 toothed. Fruit ribbed. Leaves (3)5–7 foliolate; flowers greenish white to creamy white 10

10a. Calyx deeply 5-dentate. Corolla greenish white; petal lobes distinct; stamens inserted; ovary stalked,longitudinaly ridged. Fruit ovate with longitudinal ridges, flattened compressed. Seeds completelysurrounded by hyaline wing. 1 sp. Continental South America . . . . . . . . . . . . . . . . . . . . . . . . Cybistax

10b. Calyx shallowly 5-dentate. Corolla greenish-white or white; corolla tube elongate; petal lobes united intolaciniate rim; stamens exserted or subexserted; ovary sessile, smooth or with minute ridges. Fruit linear,with distinct curve in proximal end. Seeds linear, bialate. 2 spp. Cuba and Hispanola. . . . Ekmanianthe

9b. Calyx spathaceous to irregularly bilabiate. Leaves simple to 9 foliolate. Flowers various . . . . . . . . . . . . . . 1111a. Calyx and corolla yellow, of similar texture, nearly indistinguishable. Leaves 5–7 foliolate, indumentum

of scattered lepidote scales and occasionally with simple hairs. Fruits linear, oval in cross section withirregular longitudinal ridges, irregularly costate. 2 spp. Central America to Venezuela, dry or seasonallydry areas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Roseodendron

11b. Calyx and corolla of different textures, distinguishable,although sometimes of similar color. Corollawhite to red, rarely yellow. Leaves simple to 9-foliolate, indumentum of lepidote scales, these occasionallystalked and dense, forming a tomentum. Fruits linear, without large ridges and costae described above. 67spp. Continental America to Antilles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tabebuia

ACKNOWLEDGEMENTS. This study is part of the doctoraldissertation of the first author. Thanks to Michelle Zjhra, LuciaLohmann, and one anonymous reviewer for comments thatsignificantly improved this manuscript and to Daisy Castillo(JBSD), Theodoro Clase (JBSD), Alberto Veloz (JBSD), FrankAxelrod (UPRRP), Javier Francisco-Ortega (FBG/FIU), JoseGonzalez (INBio), Monica Mejıa-Chang, Carmen Galdames(STRI), Carlos Burelo Ramos (XAL), Ricardo Rueda and DaniaPaguaga (Universidad de Nicaragua, Leon) for assistance inthe field. Thanks also to Mario Blanco, Karen Redden, John L.Clark, and H. David Clarke for collecting tissue in Ecuadorand Guyana. Urs Jauch and Merran Matthews are gratefully

acknowledged for their support with the SEM. NYBG, MOand US kindly loaned specimens for this study. Funds for thisresearch were provided by NSF DEB: 0309065, OTS, theGarden Club of America, ASPT, BSA, University of Washing-ton, Department of Botany ‘‘Plant Molecular SystematicsFellowship’’, and the University of Washington Departmentof Biology ‘‘Melinda Denton Fellowship’’.

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Bignoniaceae. Geneve: Bibliotheque Universelle.DOS SANTOS, G. and R. B. MILLER. 1992. Wood anatomy of

Tecomeae. Pp. 336–358 in Flora Neotropica Monograph. 25,ed. A. H. Gentry. Bronx: New York Botanical Garden.

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———. 1972. Handroanthus (Bignoniaceae): a critique. Taxon21: 113–114.

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———. 1992. Bignoniaceae- Part II (Tribe Tecomeae). FloraNeotropica Monographs 25. Bronx: New York BotanicalGarden.

GROSE, S. O. and R. G. OLMSTEAD. 2007. Evolution ofa charismatic neotropical tree: Molecular phylogeny ofTabebuia s. l. and allied genera (Bignoniaceae). SystematicBotany 32: 650–659.

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APPENDIX 1. List of representative herbarium specimensconsulted for mophology.

Handroanthus albus (Cham.) Mattos Reitz and Klein 7037(US) Brazil, Santa Catarina, Hatschbach, G 48929 (US) Brazil,Parana. Handroanthus barbatus (E.Mey) Mattos Prance, GT20577 (US) Brazil, Amazonas, Maguire and Wurdack 34873(US) Venezuela, Amazonas. Handroanthus billbergii (Bur &K. Schum) Grose subsp ampla Steyermark, J. 94543 (US)Venezuela, Falcon Samaniego, V 015 (US) Ecuador Loja.Handroanthus capitatus (Bur. & K. Schum) Mattos Balee, W5097 (MO) Bolivia, Beni, Blanco, C 496 (US) Venezuela, DeltaAmacuro. Handroanthus chrysanthus (Jacq.) Grose Gentry, A

1022 (MO) Ecuador, Gentry, A 14750 (US) Venezuela, DistritoFederal. Handroanthus chrysotrichus (Mart ex. DC) MattosKeisling 7228 (MO) Argentina, Buenos Aires, Peixoto, A 309(MO) Brazil, Espirito Santo. Handroanthus coralibe (Standl.)Grose Dugand, A 390b (US) Colombia, Atlantico. Han-droanthus guayacan (Seem.) Grose Gentry, A 8574 (US)Belize, Cayo, Nee, M 10447 (US) Panama, Canal Zone.Handroanthus heptaphyllus (Vell.) Mattos Pedersen, TM6526 (US) Paraguay, Misiones, Gentry, A 51909 (US) Para-guay, Paraguari. Handroanthus impetiginosus (Mart ex. DC)Mattos Killeen, T 4201 (MO) Bolivia, Santa Cruz, Nicher 35464(Z) New Caledonia, SEM. Handroanthus obscurus (Bur. & K.Schum.) Mattos Schultes, RE 16993 (US) Colombia, AmazonasPlowman, T 2758 (US) Peru, Loreto. Handroanthus ochraceus(Cham.) Mattos subsp. ochraceus Herzog 138 (Z), Bolivia,SEM. Handroanthus ochraceus (Cham.) Mattos subsp hetero-trichus Steyermark, J and Liesner, R 120985 (NY) Venezuela,Sucre. Handroanthus ochraceus (Cham.) Mattos subsp.neochrosanthus Molina, A 1379 (US) Honduras, Morazan.Handroanthus pulcherrimus (Sandwith) Grose Smith, LB 7263(US) Brazil, Santa Catarina. Handroanthus pumilus (A. H.Gentry) Grose Hatschenbach, G 49779 (US), Bazil, Han-droanthus serratifolius (Vahl) Grose Broadway, s. n. (Z)Grenada, St. Georges, SEM, Zarucchi, JL 4536 (US) Colombia,Antioquia. Handroanthus subtilis (Sprague & Sandwith)Grose Hernandez, L 469 (MO) Venezuela, Bolıvar, A. H.Gentry, A 10521 (MO) Venezuela, Bolıvar. Handroanthusuleanus (Kraenzl.) Grose Stevenson, P 955 (MO). Han-droanthus umbellatus (Sond.) Mattos A. H. Gentry, A 59133(NY) Brazil. Minas Gerais, Inacio, E 139 (MO) Brazil,Pernambuco. Handroanthus vellosoi (Toledo) Mattos A. H.Gentry, A 49602 (MO) Brazil, Minas Gerais. Roseodendronchryseum (Blake) Miranda Aristequieta 6801 (US) Venezuela.Roseodendron donnell-smithii (Rose) Miranda Bullock, SH1311 (MO) Mexico, Jalisco, Blanco, C 643 (US) Venezuela,Bolivar, S. Grose 165 (WTU), Mexico, Chiapas, SEM. Tabebuiaaurea (Manso) Benth and Hook. f.ex. S Moore Foster, P 163(MO) Bolivia, Santa Cruz, Thomas, W et al 4646 (US) Brazil,Matto Grosso. Tabebuia cristata A. H. Gentry Peixoto, A 3067(US) Brazil, Espirito Santo. Tabebuia insignis (Miq.) Sand-with var. orinocensis Maguire, B. 37701 (US) Venezuela,Amazonas. Tabebuia insignis (Miq.) Sandwith var. insignisHenkel, T 723 (US) Guyana, Potaro-Siparuni. Tabebuiainsignis (Miq) Sandw. var. monophylla Gillespie, L. J. 927(US) Guyana, Potaro-Siparuni. Tabebuia microphylla (Lam.)Urb. Zanoni, T 20808 (NY) Dominican Republic, Pedernales,Zanoni, T 36525 (MO) Dominican Republic, Pedernales.Tabebuia myrtifolia (Greenm.) A. H. Gentry var. petrophilaLeon 11489 (NY) Cuba, Havana. Tabebuia myrtifolia(Greenm.) A. H. Gentry var. myrtifolia Garcia, R 510 (MO)Dominican Republic, Pedernales. Tabebuia nodosa (Griseb.)Griseb. Hatschenbach G 49232 (US) Brazil, Curitiba. Tabebuiaophiolitica Alain A. H. Gentry A 50633 (MO) RepublicaDominica, Puerto Plata. Tabebuia orinocensis (Sandwith) A.H. Gentry Gentry, A. 14609 (US) Venezuela, Amazonas,Davidse, G. 15160 (US) Venezuela, Atures. Tabebuia pallida(Lindl.) Miers Nicholson, DH 1987 (US) Dominica, Wilbur, RL7518 (US) Dominica. Tabebuia palustris Hemsl. A. H. Gentry,A 7376 (MO) Colombia, Choco. Tabebuia pilosa A. H. GentryCuello, N 723 (US) Venezuela, Bolivar, Wurdack, JJ 39990 (US)Venezuela, Bolivar. Tabebuia rigida Urban Axelrod, F 4107(US) Puerto Rico, Puerto Rico. Tabebuia rosea (Bertol.)DCMorton, CV 2691 (US) Mexico, Oaxaca, Breteler, FJ 4418 (US)Venezuela, Zulia, S. Grose 156 (WTU), Mexico, Veracruz,SEM. Tabebuia roseo-alba (Ridley) Sandwith Prance, GT,Lleras, E, and Coelho, DF 19034 (NY) Brazil, Mato Grosso.Tabebuia stenocalyx Sprague & Stapf de Bruijn 1633 (US)Venezuela, Bolıvar.

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