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Home , Amphitecna, Bignonia, Bignoniaceae, Crescentia, Cybistax, Handroanthus

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

Taxonomic Revisions in the Polyphyletic s. l. ()

SUSAN O. GROSE1 and R. G. OLMSTEAD Department of Biology, University of Washington, Box 355325, Seattle, Washington 98195 U.S.A. 1Author for correspondence ([email protected])

Communicating Editor: James F. Smith

ABSTRACT. Recent molecular studies have shown Tabebuia to be polyphyletic, thus necessitating taxonomic revision. These revisions are made here by resurrecting two genera to contain segregate of Tabebuia. consists of the two with spathaceous calices of similar texture to the corolla. Mattos comprises the principally flowered species with an indumentum of hairs covering the and calyx. The species of Handroanthus are also characterized by having extremely dense containing copious quantities of lapachol. Tabebuia is restricted to those species with white to red or rarely yellow and having 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. chrysanthus subsp. 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, .

Tabebuia, as currently circumscribed with 100 delimitation of Tabebuia has continued into recent species, is the largest genus in Bignoniaceae and is times (Mattos 1970; Gentry 1972). distributed from the southwestern U.S. to northern Mattos (1970) divided Tabebuia into two different and . Over the course of its groups, indicating that a group of Brazilian taxa taxonomic history, it has been split and re- known as ‘‘iˆpes’’ should not remain in Tabebuia. assembled several times, as researchers interpreted His reasoning appears to have been based on the the morphological diversity in different ways. The original concept for Tabebuia, which he defined as wide range of morphological diversity suggests having simple leaves and an that in cross there may be more than one lineage included section has 3–4 /. This is within the traditional concept of Tabebuia. Recent consistent with the of Tabebuia, T. cassinoides, studies (Spangler and Olmstead 1999; Grose and but is the most exclusive concept for Tabebuia since Olmstead 2007) have confirmed that Tabebuia,as it was proposed by de Candolle (1838). Neither did currently delimited, is polyphyletic. Therefore, the Mattos think theˆ ipes belonged in , due to generic boundaries need to be redefined in their palmately compound leaves. He created the Tabebuia and related genera. The goal of this paper genus Handroanthus for those species with pal- is to revise the generic classification to be consis- mately compound leaves and 8–9 series of ovules/ tent with its phylogeny. locule, and typified this genus with Handroanthus The name Tabebuia has a long and convoluted albus (Tabebuia alba). history (Gentry 1969). This name was first published Gentry (1972, 1992) included Handroanthus in by de Candolle (Candolle 1838) who applied it to Tabebuia, insisting that Tabebuia as previously de- bignoniaceous with ‘‘simple’’ leaves. Howev- lineated is natural. Gentry (1972) was adamant that er, the concept of Tabebuia came to be expanded to the large genus Tabebuia should not be broken up encompass a large amount of morphological di- further, and expressed his ‘‘sincere hope that future versity. As researchers examined and monographed students of Bignoniaceae will consider the case of this , a number of concepts of Tabebuia Handroanthus before succumbing to further parox- emerged, producing a labyrinthine synomomy ysms of unwarranted splitting’’ (Gentry 1972). (Candolle 1838; Raffinsque 1838; Sprague and Gentry (1992) recognized some groups within Sandwith 1932). In addition, there was confusion Tabebuia, however. In his treatment for Flora as to the boundaries between Tecoma and Tabebuia Neotropica he defined 10 species groups although (Miers 1863; Seeman 1863; Bureau 1864; Schumann he placed two species, T. arimaoensis and T. 1894; Rheder 1913). Much of the confusion between heterophylla, in both groups 9 and 10. The largest Tecoma and Tabebuia was sorted out by Britton of Gentry’s groups are primarily species endemic (Britton 1915). However, disagreement as to the to the Greater Antilles (55 species). He did not use 660 2007] GROSE & OLMSTEAD: REVISION OF TABEBUIA 661 phylogenetic terminology to describe the relation- (Fig. 1C, F). The lepidote scales in a group of ships, but he clearly believed that this group of species (such as Tabebuia pilosa) may be stalked. Anillean species was monophyletic and sister to The wood anatomy of Tabebuia group I is distinc- a group of mainland species that are spread tive; it is lightweight, with medium specific gravity throughout the continental part of the genus’ range of 0.4–0.74, and lacks lapachol (dos Santos and (Gentry 1992). Miller 1992). The heartwood is indistinct from the Tabebuia donnell-smithii was placed in Tabebuia by sapwood and the rays are 1–2 cells in width (dos Rose (1892) with some reservation: ‘‘The species, Santos and Miller 1992). This group has leaves are while not agreeing in all respects with Tabebuia, ‘simple’ (or unifoliolate) or 3–7(9)-merous, depend- answers better to this than to any other known ing on species (Gentry 1992). The are linear- genus. In its and ribbed pods it is cylindrical and smooth on the surface (Gentry more like or but does not agree 1992). The flowers of Tabebuia group I are white to in other particulars.’’ This species was later moved red in color, and often have a yellow throat (Fig 1I). to Cybistax by Seibert (1940). However, Miranda Two species belonging to this , (1965) transferred this species to a new genus, and the unsampled Tabebuia nodosa have yellow Roseodendron, segregating it from Cybistax on the flowers but otherwise share the above character- basis of calyx texture and shape, sessile of istics. its ovary, and indumentum of branched hairs. Tabebuia group II can be distinguished by having Grose and Olmstead (2007) conducted a molecu- an indumentum of simple to stellate or dendroid lar phylogenetic study of the Tabebuia , hairs (Fig. 1A, D). Lepidote scales are also usually those Neotropical Bignoniaceae with palmately- present, but may be obscured by the hairs. The compound leaves. That study included 25% of all calyx is campanulate to cupular and 5-dentate Tabebuia species and had representatives from all (Fig. 1D). The wood is among the heaviest and 10 of Gentry’s (1992) species groups. The findings hardest known, with a basic specific gravity showed that Tabebuia consists of three clades. greater than 0.74 (dos Santos and Miller 1992). It ‘‘Tabebuia goup I’’ is sister to the Caribbean is also distinctive among Tabebuia s. l., in having endemic Ekmanianthe.‘‘Tabebuia group II’’ is sister very large pits (ranging from 7–14 mm) in their to Crescentieae+Spirotecoma. The third clade is vessel elements (dos Santos and Miller 1992). Only represented by T. donnell-smithii and is sister to two species in this group, T. pumila and T. Tabebuia group II +Crescentieae+ Spirotecoma. selachidentata, are described as occasionally having unifoliolate leaves (Gentry 1992). In all other cases, MATERIALS AND METHODS the leaves are digitately 3–9 foliolate. The fruits in this group are linear and slightly costate to smooth, Characters to support the phylogeny of Grose and Olmstead (2007) are based on herbarium holdings at US, and often are densely tomentose. The flowers in MO and NY and published morphological and anatomical Tabebuia group II are generally yellow (Fig. 1G) or, studies (Gentry 1992; dos Santos and Miller 1992). Herbarium in four species, T. selachidentata, T. barbata, T. specimens were consulted for morphological characters heptaphylla and T. impetiginosa, magenta with (Appendix 1). surfaces and calices were examined with a scanning electron microscope (SEM). Specimens were yellow throat (Gentry 1992). prepared for the SEM according to the protocol established Tabebuia donnell-smithii represents a group dis- in Matthews and Endress (2004) tinguished by its unique calyx (Fig. 1E). It is spathaceous in shape, and of the same color and MORPHOLOGICAL CHARACTERS DEFINING CLADES texture as the corolla. This lineage contains only one other species, T. chrysea. The indumentum is of Each of the three clades of Tabebuia: Tabebuia lepidote scales and small, glandular hairs (Fig. 1B). groups I and II and T. donnell-smithii is supported The wood is similar to that of Tabebuia group I but by several morphological characters. Gentry de- contains tyloses, or intrusive growths of the cell scribed the importance of the calyx in Bignoniaceae wall into the vessel cavities, and pits intermediate taxonomy especially in Bignonieae (Gentry in size between those of Tabebuia groups I and II 1980). This study confirms previous observations (dos Santos and Miller 1992). The corolla is yellow that calyx morphology as well as indumentum, (Fig. 1H), occasionally with thin red lines, and wood anatomy, color, and morphology fruits are linear and irregularly costate and bullate. have been important in defining the three groups of Tabebuia in this clade (dos Santos and Miller 1992). TAXONOMIC CONSEQUENCES OF THE PHYLOGENY Tabebuia group I shares the characters of an The three clades of Tabebuia s. l. correspond indumentum of lepidote scales and a spathaceous, closely to the aggregations of species groups of sometimes irregularly bi- or trilabiate calyx Tabebuia discussed by Gentry (1992), and are 662 SYSTEMATIC BOTANY [Volume 32

FIG. 1. Characters defining clades of Tabebuia s. l. Photographs are arranged above the taxon they represent as indicated in bold on the . 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. , calyx showing lepidote scales. Scale bars for each photograph as indicated. D–F. Calyx shapes and flower color. D. 2007] GROSE & OLMSTEAD: REVISION OF TABEBUIA 663 recognized as genera here. The type of Tabebuia, T. from sapwood, olive brown to blackish, lapachol cassinoides (Lamarck) A.P. de Candolle, was not present in large quantities; Rays 1–3 cells wide, included in the molecular study due to the intervessel pits large (8–14 mm), fibers thick walled; unavailability of fresh material and the lack of wood, very dense with high specific gravity (, success in extracting DNA from herbarium speci- 7.4). Leaves (3)5–9 foliolate (reported to be occa- mens. However, it has simple leaves and an sionally 1-foliolate in H. pumilus and H. selachiden- indumentum of lepidote scales covering the leaves tatus), leaflets narrowly elliptic to broadly ovate and calyx, which places it confidently in Tabebuia (17 cm long and 18.5 cm wide), with simple, group I. We now restrict Tabebuia to those species forked, stellate, barbate or dendroid at occurringinthisclade.Thiscorrespondsto least along vein axils and sometimes densely Gentry’s (1992) informal groups 2 and 6–10 of covering leaves; petiolule to 9 cm; to Tabebuia. The sister taxon to this clade, Ekma- 30 cm. Inflorescence dichotomously branched, nianthe, shares the characters described for Tabebuia without a well developed central rachis, sometimes group I. It has 9-foliolate leaves, lepidote scales, contracted; pubescence with simple, stellate, bar- and has similar wood anatomy; nevertheless, it is bate or dendroid trichomes. Flowers: Calyx co- morphologically distinct by virtue of its hawk- riaceous, campanulate 5-dentate, 4–20 mm long, 3– moth-pollinated flowers having long corolla tubes 20 mm wide; trichomes simple, stellate or den- with united lobes, and basally curved fruit. The droid, sometimes forming a dense covering. Co- autapomorphy for Tabebuia I exclusive of Ekma- rolla yellow or magenta with yellow throat, nianthe is its straight, cylindrical fruit. tubular-infundibuliform to tubular-campanulate, The second clade, Tabebuia group II, comprises tube 2.5–6.5 cm long, 0.6–3.5 cm wide at mouth, most of the species with yellow-flowers, and lobes 0.5–5 cm; tube glabrous to densely tomentose includes many taxa segregated by Mattos (1970) with stellate to dendroid or barbate trichomes; into Handroanthus as well as Gentry’s (1992) lobes sometimes pilose in sinuses; throat papillose, informal groups 3–5. The type of Handroanthus, pubescent at level of insertion. H. albus (Tabebuia alba), is not included in this study didynamous; thecae divaricate, 1.5–2 mm (20 mm due to lack of available tissue, but it shares the four in H. subtilis) reduced. Ovary conical to characters listed above and is assignable unambig- linear-oblong; ovules 2–10 seriate in each locule. uously to this clade. Gentry (1972) insisted that Fruit an elongate linear to cylindric , Couralia had nomenclatural priority over Han- smooth to slightly costate, glabrous to scattered droanthus as circumscribed by Mattos (1970). lepidote to pubescent; pubescence scattered to However, the type of Couralia, Tabebuia fluviatilis villous; trichomes simple, stellate or dendroid. (Aubl) A.P. de Candolle, is assigned here to thin, bialate; wings hyaline membranacous Tabebuia I, so the name Couralia remains a and sharply demarcated from body. for Tabebuia. Accordingly, priority rests with A genus of 30 species distributed throughout Handroanthus Mattos (Mattos 1970). Central and with one species (H. Tabebuia donnell-smithii was placed by Seibert billbergii) in the Antilles. (1940) in the genus Cybistax and later in Roseoden- Taxa transferred to Handroanthus (*indicates the dron by Miranda (1965). Roseodendron is resurrected nine taxa used in Grose and Olmstead 2007): for T. donnell-smithii and T. chrysea, since the latter, like T. donnell-smithii has calyces that are texturally 1. (Chamisso) Mattos, Loef- identical to the corolla. These two species form grenia 50: 2. 1970. Tecoma alba Chamisso, Gentry’s (1992) informal group 1. Linnaea 7: 655. 1832. Tabebuia alba (Chamisso) Sandwith, Lilloa 14: 136. 1948.—TYPE: BRA- TAXONOMIC DESCRIPTIONS ZIL. Parana´ and , Sellow s.n. (B, lectotype HBG). HANDROANTHUS Mattos, Loefgrenia 50: 2 1970.— 2. H. arianeae (A. H. Gentry) S. Grose, comb. nov. TYPE: Handroanthus albus (Cham.) Mattos. Tabebuia arianae Gentry, Flora Neotropica Trees or occasionally ; heartwood distinct 25(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 with yellow 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). 664 SYSTEMATIC BOTANY [Volume 32

3. H. BARBATUS (E. Mey.) Mattos, Loefgrenia 50: 2. a. H. chrysanthus subsp. chrysanthus* 1970. barbata E. Mey. Nova Acta Tabebuia rufescens J. R. Johnston, Proc. Amer. Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Acad. Arts 40: 696. 1905. Cur. 12: 782. 1825. barbata (E. Meyer) Tecoma evenia Donn. Sm., Bot. Gaz. (Crawfords Miquel. Flora 25: 430. 1842. Tecoma barbata (E. ville) 20. 1895 pro parte. Meyer) DC. Prodr. 9: 221. 1845. Tabebuia Tecoma palmeri Kraenzl. Feddes Repert, 17: 220. barbata (E. Meyer) Sandwith, Lilloa 3: 462. 1921. 1938. b. H. chrysanthus subsp. meridionalis (A. H. Bignonia fluviatilis Aublet Humboldt, Bon- Gentry) S. Grose, comb. nov. Tabebuia pland and Kunth, Nov. gen. sp. Pl3: 139. 1819 chrysantha ssp. meridionalis A. Gentry, Phy- non Aublet. tologia 35: 193. 1977. Tecoma toxophora Martius emend DC, Prodr. 9: 217. Tecoma spectabilis Planchon and Linden Fl. Serres 1845 (excl. sin. Margr.) non Martius, Flora 24, Jard. L’Europe. 9: 233. 1854. Beibl. 15. 1841. Couralia toxophora (Martius Tabebuia spectabilis (Planch. and Linden) Ni emend DC) Bentham and Hooker f. ex K. chols., Ill. dict. gard. 4: 1. 1887. Schum., Nat. Pflanzenfam. 4(3b): 239. 1894. c. H. chrysanthus subsp. pluvicolus (A. H. 4. H. billbergii (Bur. & K.Schum.) S. Grose, comb. Gentry) S. Grose comb. nov.* Tabebuia nov. Tecoma billbergii Buerau & K. Schum. in chrysantha (Jacquin) Nicholson subsp. pluvi- Martius, Fl. Bras. 8(2): 319. 1897. cola. A. Gentry in Phytologia 35: 190. 1977. a. H. billbergii. subsp. billbergii. Tabebuia bill- Tecoma grandis Appun., Behand. Samereien und bergii subsp. billbergii Bureau & K. Schum. in Pflanzen 39. 1858, nom. nud. Martius, Fl. Bras 8(2): 319. 1897. 10. H. CHRYSOTRICHUS (Mart. ex DC) Mattos, b. H. billbergii subsp. ampla (A. H. Gentry) S. Loefgrenia 50: 2. 1970.* Tecoma chrysotricha Grose, comb. nov. Tabebuia billbergii subsp. Mart. ex. DC, Prodr. 9: 216. 1845. Gelseminum ampla A. Gentry, Phytologia 35: 187. 1977. chrysotrichum (Mart. ex. DC) O. Kuntze, Rev. Tabebuia ecuadoriensis Standley, Trop. 46: gen. pl. 3(2): 245. 1898. 17. 1936. Tecoma ochracea var. denudata Chamisso, Linnaea 7: 5. H. botelhensis (A. H. Gentry) S. Grose, comb. 653. 1832. nov. Tabebuia botelhensis A. Gentry, Flora Tecoma flavescens Mart. ex. DC, Prodr. 9: 216. 1845. Neotropica 25(2): 150. 1992. Tecoma obstusata DC, Prodr. 9: 217. 1845. 6. H. bureavii (Sandwith) S. Grose, comb. nov. Tecoma chrysotricha var. obtusata (DC) Bur. & K. Tabebuia bureavii Sandwith, Kew Bull. 1958: Schum, Martius Fl. Bras. 8(2): 338. 1897. 442. 1992. Tecoma dentata Bur. & K. Schum, Tabebuia chrysotricha (Mart. ex. DC Standley, Martius Fl. Bras. 8(2): 323. 1897, not Tabebuia Publ. Field. Mus. Nat. Hist. Bot. Ser. 11: 176. dentata, 1863. Handroanthus dentatus (Bur. and 1936. Tabebuia chrysothrica var. obtusata K. Schum.) Mattos, Loefgrenia 50: 2. 1970. (DC)Toledo, Arq. bot. estado Sa˜o Paulo 3(1): 7. H. CAPITATUS (Bureau ex. K. Schum.) Mattos, 35. 1952. Handroanthus chrysotrichus var. obtu- Loefgrenia 50: 2. 1970. Tabebuia capitata (Bu- sata (DC) Mattos, Loefgrenia 50: 2. 1970. reau ex K. Schum.) Sandwith, Rec. Trav. Bot. Tecoma pedicellata Bur & K. Schum., Martius Fl. Neerl. 34: 226. 1937. Tecoma capitata Bureau & Bras. 8(2): 336. 1897. Handroanthus pedicellatus K. Schum., Fl. Bras. 8(2): 337. 1897. (Bureau & K. Schum.) Mattos, Loefgrenia 50: 2. Tecoma leucoxylon (L.) Mart. ex. DC var. miquelii 1970. DC, Prodr. 9: 219. 1845. Tecoma grandis Kraenzl., Feddes Repert. 17: 217. Tabebuia glomerata Urban, Feddes Rebert. 14: 305. 1921 1916. 11. H. coralibe (Standley) S. Grose comb. nov. Tabebuia hypolepra Sprague and Sandwith, Kew Tabebuia coralibe Standley in Trop. Woods. 36: Bull. 1932: 25 1932. 18. 1933. 8. H. catarinensis (Gentry) S. Grose comb. nov. 12. H. cristatus (Gentry) S. Grose comb. nov. Tabebuia catarinensis Gentry, Ann. Missouri Tabebuia cristata A. Gentry in Flora Neotropica Bot. Gard. 64: 318. 1977. 25(2): 174. 1992. 9. H. chrysanthus (Jacq.) S. Grose comb. nov. 13. H. guayacan (Seeman) S. Grose comb. nov.* Bignonia chrysantha Jacq., Pl. hort. Schoenbr. Tecoma guayacan Seemann, Bot. voy. Herald 2: 45, tab. 211. 1797. Tecoma chrysantha (Jacq) 180. 1854. Tabebuia guayacan (Seemann) Hems- DC, Prodr. 9: 211. 1845. Tabebuia chrysantha ley in Biol. Centr.-Amer Bot. 2: 495. 1882. (Jacq.) Nichols. subsp. Chrysantha, Ill. dict. 14. H. HEPTAPHYLLUS (Vell.) Mattos, Loefgrenia 50: gard. 4: 1. 1887. 2. 1970. Bignonia heptaphylla Vell., Fl. Flumin. 2007] GROSE & OLMSTEAD: REVISION OF TABEBUIA 665

251. 1829. Tabebuia heptaphylla (Vell.) Toledo, Tabebuia schunkevigoi Simpson, Fieldiana, Bot. 36: 1. Arq. Bot. Estado Sa˜o Paulo, n. s. 3: 33. 1952. 1972. Tecoma eximia Miq., Linnaea 22: 803. 1849. Tabebuia 16. H. incanus (A. H. Gentry) S. Grose comb. nov. eximia (Miq.) Sandwith, Lloydia 2: 213. 1939. Tabebuia incana A. Gentry in Ann. Missouri Handroanthus eximia (Miq.) Mattos, Loefgrenia Bot. Gard. 65: 732. 1978. 50: 2. 1970. 17. H. lapacho (K. Schum.) S. Grose comb. nov. Tecoma curialis Saldanha da Gama, Config, descr. Tecoma lapacho K. Schum., Pflanzenfam. 4(3b): orga˜os fund. de Janeiro. 1: 51. 1865. 238. 1894. (K. Schum.) Sand- Tecoma ipe Mart. ex. K. Schum., Engl. and Prantl, with, Lilloa 14: 136. 1948. Pflanzenfam. 4(3b): 238. 1894. Tabebuia ipe Tabebuia flavescens (Vell.) Griseb., sensu Griseb., (Mart. ex. K. Schum.) Standley, Trop. Woods. Symb. fl. Argent. 257. 1879. 36: 20. 1933. 18. H. OBSCURUS (Bur. ex. K Schum.) Mattos, Tecoma ipe var. desinens Sprague, Bull. Herb. Loefgrenia 50: 2. 1970.* Tecoma obscura Bur. Boissier, ser. 2, 5: 86. 1905. ex K. Schum., Mart, Fl. Bras. 8(2): 343. 1897. Tecoma ipe var. desinens f. parviflora Sprague, Bull. Tabebuia obscura (Bureau ex. K. Schum.) Sand- Herb. Boissier, ser. 2, 5: 86. 1905. with, Rec. Trav. Bot. Neerl. 34: 226. 1937. Tecoma ipe var. desinens f. grandiflora Sprague, Bull. Tabebuia subtilis var. schultesiana Sandwith, Bot. Herb. Boissier, ser. 2, 5: 86. 1905. Mus. Leafl. Harvard Univ. 17: 96. 1955. Tabebuia avellanedae var. paulensis Toledo, Arq. Bot. Tabebuia obscura var schultesiana (Sandwith) Estado Sa˜o Paulo, n. s., 3: 1952. Sandwith in Dugand, Mutisia 25: 16. 1956. Tabebuia impetiginosa var. lepidota (Bur.) Toledo, 19. H. OCHRACEUS (Cham.) Mattos, Loefgrenia 50: Arq. Bot. Estado Sa˜o Paulo, n. s., 3: 1952. 2. 1970. Handroanthus impetiginosus var. lepidotus (Bur.) a. H. OCHRACEUS subsp. HETEROTRICHUS (DC) S. Mattos, Loefgrenia 50: 2. 1970. Grose. Tecoma heterotricha DC, Prodr. 9: 219. 15. H. IMPETIGINOSUS (Mart. Ex DC) Mattos, Loef- 1845. Tabebuia heterotricha (DC) Hemsley, grenia 50: 2. 1970.* Tecoma impetiginosa Mart. Biol. Centr.-Amer., Bot. 2: 495. 1882. Tabe- ex. DC, Prodr. 9: 218. 1845. Tabebuia impetigi- buia ochracea (Cham) Standley subsp. hetero- nosa (Mart. ex. DC) Standley, Publ. Field Mus. trichus (DC) A. H. Gentry, Fl. Nat. Hist., Bot. Ser. 11: 176. 1936. 8(4): 391. 1982. Tabebuia avellanedae Lorentz ex. Griseb., Symbol. fl. b. H. OCHRACEUS subsp. NEOCHRYSANTHUS (A. H. argent. 258. 1879. Gelseminum avellanedae (Lor- Gentry) S. Grose.* Tabebuia neochrysantha A. entz ex Grisebach) Kuntze, Rev. gen. pl. 3(2): H. Gentry, Brittonia 22: 260. 1970. Tabebuia 245. 1898. Handroanthus avellanedae (Lorentz ex ochracea (Chamisso) Standley subsp. neo- Griseback) Mattos, Loefgrenia 50: 2. 1970. chrysantha (A. H. Gentry) A. H. Gentry, Tabebuia palmeri Rose, Contr. U. S. Natl. Herb. 1: Ann. Missouri Bot. Gard. 60: 948. 1974. 109. 1891. Tabebuia chrysantha (Jacq.) Nichols., Dict. Gard. 4: 1. Tecoma impetiginosa var lepidota Bureau, Vidensk. 1897, sensu Sandwith, non Jacq. Meddel. Dansk. Naturhist. Foren. Kjøbenhavn. c. H. OCHRACEUS (Cham.) Mattos subsp. OCHRA- 1893: 114. 1894. Handroanthus impetiginosus CEUS. Tecoma ochracea Cham., Linnaea 7: 653. var. lepidotus (Bureau) Mattos, Loefgrenia 50: 1832. 2. 1970. Tecoma hypodictyon DC, Prodr. 9: 217. 1845. Tabebuia Tecoma adenophylla K. Schum. ex. Bureau & K. hypodiction (DC) Standl., Field Mus. Nat. Hist., Schum. in Martius, Fl. Bras. 8(2): 412. 1897. Bot. Ser. 11: 176. 1936. Tecoma ipe var. integra Sprague, Bull. Herb. Biossier, Tecoma heteropoda DC, Prodr. 9: 219. 1845. Tabebuia ser. 2 5: 86. 1905. Tecoma integrum (Sprague) ochracea subsp. heteropoda (DC) A. H. Gentry, Chodat, Bull. Soc. Bot. Gene`ve ser. 2, 9: 242. in Prance, Biological Diversification in the 1917. Tabebuia ipe var. integra (Sprague) Sand- 132. 1982. with, Lloydia 2: 213. 1939. Bignonia tomentosa Pav. ex DC, Prodr. 9: 219. 1845. Tecoma ipe var. integrifolia Hassler, Rev. Inst. Parag. nom. nud., pro syn. 3: 166. 1901. Tecoma hassleri Sprague, Trans. Proc. Bot. Soc. Tecoma avellanedae var. alba Lillo, Seg. contr. arb. Edinburgh 48: 435. 1904. Argent. 13. 1917. Tecoma grandiceps Kra¨nzl., Feddes Repert. 17: 216. 1921. Tabebuia nicaraguensis Blake, Contr. Gray Herb. 52: Tecoma campinae Kraenzlin, Feddes Repert. 17: 215. 95. 1917. 1921. Tabebuia dugandii Standley, Trop. Woods 36: 17. Tecoma hemmendorffiana Kraenzl., Feddes Repert. 1933. 17: 224. 1921. 666 SYSTEMATIC BOTANY [Volume 32

20. H. PEDICELLATUS (Bur. ex K. Schum.) Mattos, comb. nov.* Tabebuia subtilis Sprague and Loefgrenia 50: 2. 1970. Tecoma pedicellata Bur. & Sandwith, Kew Bull. 1932: 23. 1932. K. Schum., in Mart., Fl. Bras. 8(2): 336. 1897. 28. H. uleanus (Kraenzl) S. Grose comb. nov. Tabebuia pedicellata (Bur. ex. K. Schum.) A. H. Tecoma uleana Kra¨nzlin, Feddes Repert. 17: Gentry, Flora Neotropica 25(2): 236. 1992. 217. 1921. Tabebuia uleana (Kraenzl.) A. H. Tecoma catinga Bur. & K. Schum., in Martius, Fl. Gentry in Mem. N.Y. Bot. Gard. 29: 279. 1978. Bras. 8(2): 337. 1897. Handroanthus catinga (Bur. 29. H. UMBELLATUS (Sonder) Mattos, Loefgrenia 50: & K. Schum.) Mattos. Loefgrenia 50: 4. 1970. 2. 1970.* Tecoma umbellata Sond., Linnaea 22: 21. H. pulcherrimus (Sandwith) S. Grose comb. 562. 1849. Tabebuia umbellata (Sond.) Sandwith, nov. Tabebuia pulcherrima Sandwith, Lilloa 14: Lilloa 14: 136. 1948. 133. 1948. Tecoma petropolitana Glaz. nom. Tecoma eximia Miq., Linnaea 22: 803. 1849. Tabebuia nud., Mem. Soc. Bot. France 3: 528. 1911. eximia (Miq.) Sandwith, Lloydia 2: 213. 1939. 22. H. pumilus (A. H. Gentry) S. Grose comb. nov. Handroanthus eximius (Miq.) Mattos, Loefgre- Tabebuia pumila A. H. Gentry, Flora Neotropica nia 50: 2. 1970. 25(2): 244. 1992. Tecoma umbellata var. lanceolata Bur. & K. Schum., 23. H. riodocensis (A. H. Gentry) S. Grose comb. Martius, Fl. Bras. 8(2): 335, 1897. Tabebuia um- nov. Tabebuia riodocensis A. H. Gentry, Flora bellata var. lanceolata (Bur. & K. Schum.) Toledo, Neotropica 25(2): 1248. 1992. Arq. Bot. Estado Sa˜o Paulo 3(1): 35. 1952. 24. H. selachidentatus (A. H. Gentry) S. Grose Handroanthus umbellatus var. lanceolatus (Bur. & comb. nov. Tabebuia selachidentata A. H. Gen- K. Schum.) Mattos, Loefgrenia 50: 2. 1970. try, Flora Neotropica 25(2): 254. 1992. 30. H. VELLOSOI (Toledo) Mattos, Loefgrenia 50: 2. 25. H. serratifolius (Vahl) S. Grose comb. nov.* 1970. Tabebuia vellosoi Toledo, Arq. Bot. Estado Bignonia serratifolia Vahl, Eclog. Amer. 2: 46. Sa˜o Paulo, n. s. 3(1): 34. 1952. 1798. Tecoma serratifolia (Vahl) G. Don., Gen. Bignonia longiflora Vell., Fl. flumin. 252. 1829 (1825). syst. 4: 224. 1838. Tabebuia serratifolia (Vahl) Tecoma longiflora (Vell.) Bur. & K. Schum., in Nichols. in Dict. Gard. 4: 1. 1887. Martius, Fl. Bras. 8(2): 324. 1897, non Tecoma Bignonia flavescens Vell., Fl. flumin. 252. 1829 (1825). longiflora Griseb. 1866. Tecoma flavescens (Vell.) Martius ex DC, Prodr. Tecoma alba Cham. var. subdenudata Bur., Dansk. 9: 226. 1845. Handroanthus flavescens (Vell.) Naturhist. Foren. Kjøbenhavn 1893: 115. 1893. Mattos, Loefgrenia 50: 2. 1970. Bignonia araliacea Cham., Linnaea 7: 683. 1832. ROSEODENDRON Miranda, Bol. Soc. Not. Mex. 29:43 Tecoma araliacea (Cham.) DC, Prodr. 9: 221. 1965.—TYPE: Roseodendron donnell-smithii 1845. Tabebuia araliacea (Cham.) Morong and (Rose) Miranda. Britton, Ann. N. Y. Acad. Sci. 7: 190. 1893. Trees to 35 m; heartwood not very distinct from Gelseminum araliaceum (Cham.) O. Kuntze, sapwood, yellowish or light to reddish brown; Rev. gen. 3(2): 245. 1898. Handroanthus aralia- tyloses present; lapachol present in small quanti- ceus (Cham.) Mattos, Loefgrenia 50: 2. 1970. ties (R. chrysea); rays 2–5(8) cells wide; medium Tecoma conspicua DC, Prodr. 9: 221. 1845. Bignonia intervessel pits (6–8 mm); thin to thick walled conspicua Richard ex DC, Prodr. 9: 221, nom. fibers; medium specific gravity (0.4–7.4). Leaves nud., pro syn. palmately 5–7 foliolate; leaflets oblong-elliptic, to Tecoma patrisiana DC, Prodr. 9: 221. 1845. 28 cm long and 14 cm wide, usually much smaller, Tecoma speciosa DC, Prodr. 9: 218. 1845. Gelseminum membranaceous to chartaceous, the surface some- speciosum (DC) O. Kuntze, Rev. gen. 3(2): 245. what rough, especially above, sometimes puber- 1898. ulous above and below with simple or dendroid Tecoma nigrescens Klotz in Schomburgk, Reisen 3: trichomes, or more or less glabrate, drying olive, 1159. 1848, nom. nud. darker above and lighter below; apex acute to Tecoma atractocarpa Bur. & K. Schum., Martius, Fl. acuminate; bases truncate; petiolules to 7 cm long; Bras. 8(2): 326. 1897. petiole to 26 cm long, tomentose with dendroid Handroanthus atractocarpus (Bur. & K. Schum.) and barbate trichomes or glabrate. Inflorescence Mattos, Loefgrenia 50: 2. 1970. a large terminal panicle with the central rachis well Tabebuia monticola Pittier, Cat. Flora Venez. 2: 409. developed and lateral branches either very short 1947, nom. nud. and dendroid-pubescent, or branching with rachis 26. H. spongiosus (Rizzini) S. Grose comb. nov. and branches puberulous with capitate trichomes. Tabebuia spongiosus Rizz., Rodriguesia 28: 172. Flowers: Calyx campanulate, 10–20 mm long, 5– 1976. 15 mm wide, finely membranaceous, bilabiate to 27. H. subtilis (Sprague & Sandwith) S. Grose irregularly several–5-dentate, somewhat lepidote 2007] GROSE & OLMSTEAD: REVISION OF TABEBUIA 667 and puberulous, sometimes with gland tipped Trees or shrubs; heartwood not distinct from trichomes. Corolla yellow, sometimes with reddish sapwood, light to reddish brown; lapachol absent; lines in throat, tubular-infundibuliform; tube 3– rays 1–3 cells wide; small to medium intervessel 4.5 cm long, 0.8–2 cm wide at mouth of tube, lightly pits (3–6 mm); thin to thick walled fibers; medium puberulous to almost glabrous outside, generally specific gravity (0.4–7.4). Leaves 1–7(9) foliolate; glabrous inside, pubescent at level of stamen leaflets narrowly elliptic to orbicular to 35 cm long insertion; lobes glabrous to lightly pubescent; lobes and 32 cm wide, with stalked or sessile lepidote 1–2.5 cm long. Stamens didynamous; thecae di- scales that are usually scattered and sometimes varicate, 2–2.5 mm long; staminode reduced. Ovary densely covering undersurface of leaves; petiolule linear, 5–6 mm long, 0.6–1.5 mm wide; ovules 4 or 6 to 11 cm; petiole to 18 cm. Inflorescence dichoto- seriate in each locule. Fruit an elongate-linear mously branching, without a well developed central capsule, 25–45 cm long, 0.9–3 cm wide, longitudi- rachis, usually a few flowered panicle, occasionally nally striate-costate to irregularly striate costate with many flowered, sometimes densely lepidote. Flow- 8–12 rather irregular longitudinal ribs, puberulous ers: Calyx coriaceous, 2–3 labiate rarely 5-dentate, with simple and branched or simple-gland-tipped 5–25 mm long, 4–11 mm wide, densely lepidote. trichomes. Seeds thin; wings hyaline-membranac- Corolla white to red, often with yellow throat, eous and sharply demarcated from seed body. completely yellow in two species (T. nodosa and T. Two species ranging from Me´xico to Northwest- aurea) tubular-infundibuliform to tubular-campan- ern South America, and dry areas of Venezuela ulate; tube 2–7 cm long, 0.6–3.5 cm wide at mouth; and . lobes 0.5–3.2; tube glabrous to pubescent at level of Taxa transferred to Roseodendron (* indicates stamen insertion. Stamens didynamous; thecae taxon used in Grose and Olmstead, 2007): divaricate, 2–6 mm; staminode reduced. Ovary linear; ovules 2–3 seriate in each locule. Fruit an elongate linear to cylindric capsule, smooth to 1. ROSEODENDRON CHRYSEUM (Blake) Miranda, Bol. Soc. Mex. 29: 43. 1965. Tabebuia chrysea Blake, costate striate, minutely to densely lepidote. Seeds Contr. Gray herb. 53: 50. 1918. Tecoma chrysea thin, bialate; wings hyaline membranacous and (Blake) Pittier, publ. Pl. Usual. Venez. 63. 1939. sharply demarcated from seed body. Cybistax chrysea (Blake) Seibert, Trop Woods A genus of 67 species widely distributed 63: 7. 1940. throughout Central and South America and the Antilles. 2. R. DONNELL-SMITHII (Rose) Miranda, Bol. Soc. Included taxa (* indicates taxon used in Grose Mex. 29: 43. 1965.* Tabebuia donnell-smithii and Olmstead 2007). Names in this group remain Rose, Bot. Gaz. 17: 418. pl. 26. 1892. Cybistax unchanged from Gentry (1992) and are listed for donnell-smithii (Rose) Seibert, Carnegie Inst. convenience: Wash. Publ. 522: 392. 1940.—TYPE: GUATE- MALA. Escuintla, Donnell-Smith 2070 (US!). 1. TABEBUIA ACROPHYLLA (Urb.) Britton* Cybistax millsii Miranda, Bol. Soc. Mex. 26: 129. 2. T. ANGUSTATA Britton 1961, Roseodendron millsii (Miranda) Miranda, 3. T. ARIMAOENSIS Britton Bol. Soc. Mex. 29: 43. 1965, Tabebuia millsii 4. T. AUREA (Silva Manso) Benth. & Hook. f. ex S. (Miranda) A. H. Gentry, Ann. Missouri Bot. Moore* Gard. 63: 75. 1976. 5. T. BAHAMENSIS (Northr.) Britton* 6. T. BERTEROI (DC) Britton* TABEBUIA Gomes ex A.P. de Candolle, Biblioth. 7. T. BIBRACTEOLATA (Griseb.) Britton Universelle Gene`ve, ser 2, 17:130. Sep. 1838.— 8. T. BROOKSIANA Britton TYPE: Tabebuia uliginosa (Gomes) DC 5 9. T. BUCHII (Urb.) Britton (Lam.) DC, Encyc. Me´th. 10. T. BULLATA A. H. Gentry 1: 418. 1785. 11. T. CALCICOLA Britton Leucoxylon Raf., Sylva Tellur. 77. Oct 1838.—TYPE: 12. T. CALETICANA A. H. Gentry and D. Albert L. riparia Raf. 5 T. heterophylla (DC) Britton. 13. T. CASSINOIDES (Lam.) DC (Type) Potamoxylon Raf., Sylva Tellur. 78. Oct 1838.— 14. T. CLEMATIS Alain TYPE: P. alba Raf. 5 T. fluviatilis (Aubl. ) DC. 15. T. CONFERTA Urb. Proterpia Raf., Sylva Tellur. 80. Oct 1838.—TYPE: P. 16. T. CRISPIFLORA Alain obtusifolia (Lamarck) Raf. (Bignonia obtusifolia 17. T. XDEL-RISCOI Borhidi Lam.) 5 T. cassinoides (Lamarck) DC 18. T. DENSIFOLIA Urb. Couralia Splitgerberger, Tujdschr. Natuurl. Gesch. 19. T. DOMINGUENSIS (Urb.) Britton Physiol. 9: 14. 1842.—TYPE: Couralia fluviatilis 20. T. DUBIA (C.Wright ex Sauvalle) Britton ex (Aubl.) Splitgerberger 5 T. fluviatilis (Aubl.) DC. Seibert 668 SYSTEMATIC BOTANY [Volume 32

21. T. ELEGANS (Urb.) 50. T. PLATYANTHA (Griseb.) Britton 22. T. ELLIPTICA (DC) Sandwith 51. T. POLYANTHA Urb. & Ekman 23. T. FLUVIATILIS (Aubl.) DC 52. T. POLYMORPHA Urb. 24. T. GLAUCESCENS Urb. 53. T. PULVERULENTA Urban 25. T. HAEMANTHA (Bertero ex Sprengel) DC* 54. T. RETICULATA A. H. Gentry 26. T. HETEROPHYLLA (DC) Britton* 55. T. REVOLUTA (Urb.) Britton 27. T. HYPOLEUCA (Wright ex Sauvalle) Urb. 56. T. RIGIDA Urb. 28. T. INAEQUIPES Urb. 57. T. ROSEA (Bertol.) DC* 29. T. INSIGNIS (Miq.) Sandwith 58. T. ROSEO-ALBA (Ridl.) Sandwith 59. T. SAUVALLEI Britton* a. T. INSIGNIS (Miq.) Sandwith var. INSIGNIS* 60. T. SCHUMANNIANA Urb.* INSIGNIS MONO- b. T. (Miq.) Sandwith var. 61. T. SHAFERI Britton PHYLLA Sandwith 62. T. SIMPLICIFOLIA Carabia ex Alain c. T. INSIGNIS (Miq.) Sandwith var. PACIMO- 63. T. STENOCALYX Sprague & Stapf NENSIS Sandwith 64. T. STRIATA A. H. Gentry* 65. T. TRACHYCARPA (Griseb.) K.Schum. 30. T. JACKIANA Ekman ex Urb. 66. T. VINOSA A. H. Gentry 31. T. LEPIDOPHYLLA (A. Richard) Greenm. 67. T. ZANONII A. H. Gentry 32. T. LEPIDOTA (HBK) Britton* 33. T. LEPTONEURA Urb. Tabebuia Alliance. The character uniting the 34. T. LINEARIS Alain members of the Tabebuia alliance and distinguish- 35. T. MAXONII Urb. ing them from other Bignoniaceae is digitately- 36. T. MICROPHYLLA (Lam.) Urb.* compound leaves. Members of this alliance are 37. T. MOAENSIS Britton usually shrubs to large trees, and the leaves are 38. T. MULTINERVIS Urb. and Ekman often covered with lepidote scales and sometimes 39. T. MYRTIFOLIA (Griseb.) Britton with simple or branched hairs. The flowers have a calyx that is spathaceous or cupular with 3–5 a. T. MYRTIFOLIA (Griseb.) Britton var. MYR- lobes. The calyx and corolla often have an TIFOLIA indumentum of lepidote scales or hairs. The ovary b. T. MYRTIFOLIA (Griseb.) Britton var. PET- is linear or ovate, and bilocular at least in extreme ROPHILA (Greenm) A. H. Gentry base. The fruit has striations, ridges or spines, and is usually loculicidally dehiscent but is indehiscent 40. T. NODOSA (Griseb.) Griseb. in the tree genera of the Crescentieae. The seeds of 41. T. OBOVATA Urb. wind-dispersed taxa are winged, those that are 42. T. OBTUSIFOLIA (Cham.) Bur. water or mammal dispersed have vestigial wings. 43. T. OPHIOLITICA Alain KeytotheGenera. Not all specimens are 44. T. ORINOCENSIS (Sandwith) A. H. Gentry collected at a time of year to show all of the 45. T. PALLIDA (Lindl.) Miers desired features, usually a combination of leaves, 46. T. PALUSTRIS Hemsl.* flowers, and fruit. This key is intended to be also 47. T. PANICULATA Leonard useful for specimens that are sterile or contain only 48. T. PILOSA A. H. Gentry flowers and fruits. The most distinguishing fea- 49. T. PINETORUM Britton 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, sometimes with darker lines in throat. Fruit linear, terete capsule, valves subwoody. Seeds linear, seed wings made of minute hairs. One sp. Continental South America ...... 1b. Young branches and inflorescence not shiny with sticky secretions, instead with indumentum of hairs and/or lepidote scales. 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 stalked lepidote, 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 , covered with simple hairs, flowers small, yellow/brown occasionally with mauve in throat. Fruit linear, spirally coiled. Seeds bialate with hyaline wings. 2 spp. Dry in Central and South America ...... Godmania 3b. Leaves 3–9 foliolate, not glandular punctate, with indumentum of stellate or branched hairs, often densely covering surface. These hairs also present on new growth. Inflorescence various often with stellate hairs, especially on calyx and ovary. Flowers and fruit various ...... 4 2007] GROSE & OLMSTEAD: REVISION OF TABEBUIA 669

4a. shrubs or , to 3 m tall. Leaves 5-foliolate, with indumentum of dense stellate or dendroid tomentose 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 with stellate hairs. Fruit flattened orbicular, with spiny projections, also densely stellate tomentose. Seeds orbicular. 2 spp. Restricted to Brazilian ...... Zeyheria 4b. Plants usually trees, rarely shrubs. Leaves 3–9 foliolate, reportedly 1-foliolate in one Brazilian species, with indumentum 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, calyx usually 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 or white to red...... 5 5a. cauliflorous or ramiflorous, growing from round or branchlike areas on (terminal in some species of ) ...... 6 6a. Fruit dehiscent, linear, sometimes curved or coiling; septum thickened with pits in which seeds rest. Seeds very small. Leaves simple or 3–7 foliolate, shiny when dry, drying dark, undersurface with simple hairs forming domatia, 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 or cauliflorous; calyx tubular-campanulate, irregularly dentate; flowers with elongate; corolla yellow to dark red, lower corolla lobes reflexed; stamens exserted, 4 spp. and Hispan˜ ola ...... Spirotecoma 6b. Fruit indehiscent, linear to spherical, seeds wingless, or with vestigial wings. Leaves usually simple, when compound 3–5 foliolate, not drying as above, membranaceous, glabrous or lepidote, sometimes with stipular spines. 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 small with vestigial wings, no pulp in fruit. Flowers terminal and cauliflorous, greenish white; calyx membranaceous, irregularly spathaceously split. 8 spp. Me´xico to extreme NW Colombia . . 7b. Leaves simple (in part trifoliolate in alata). Fruit a pepo, bilocular to approximately halfway. Seeds embedded in white fleshy, sweet pulp. Inflorescences cauliflorous or terminal, calyx thick, tearing irregularly. 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, lobes united into rim. Fruits oblong (if orbicular, then mangrove species). Seeds very large, 2cmwingless. 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 lobes with long, drooping apices. Fruit, green, orbicular. Seeds very small and numerous. 6 spp. Central America and Greater Antilles, one species cultivated worldwide (C. cujete) ...... Crescentia 5b. Inflorescences terminal or axillary never cauliflorous or ramiflorous ...... 9 9a. 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 completely surrounded 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 into laciniate 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 Hispan˜ ola. . . . Ekmanianthe 9b. Calyx spathaceous to irregularly bilabiate. Leaves simple to 9 foliolate. Flowers various ...... 11 11a. 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 with irregular longitudinal ridges, irregularly costate. 2 spp. Central America to Venezuela, dry or seasonally dry areas ...... Roseodendron 11b. Calyx and corolla of different textures, distinguishable,although sometimes of similar color. Corolla white to red, rarely yellow. Leaves simple to 9-foliolate, indumentum of lepidote scales, these occasionally stalked and dense, forming a tomentum. Fruits linear, without large ridges and costae described above. 67 spp. Continental America to Antilles ...... Tabebuia

ACKNOWLEDGEMENTS. This study is part of the doctoral acknowledged for their support with the SEM. NYBG, MO dissertation of the first author. Thanks to Michelle Zjhra, Lu´cia and US kindly loaned specimens for this study. Funds for this Lohmann, and one anonymous reviewer for comments that were provided by NSF DEB: 0309065, OTS, the significantly improved this manuscript and to Daisy Castillo Garden Club of America, ASPT, BSA, University of Washing- (JBSD), Theodoro Clase (JBSD), Alberto Veloz (JBSD), Frank ton, Department of Botany ‘‘Plant Molecular Systematics Axelrod (UPRRP), Javier Francisco-Ortega (FBG/FIU), Jose´ Fellowship’’, and the University of Washington Department Gonzalez (INBio), Mo´nica Mejı´a-Chang, Carmen Galdames of Biology ‘‘Melinda Denton Fellowship’’. (STRI), Carlos Burelo Ramos (XAL), Ricardo Rueda and Dania Paguaga (Universidad de , Leo´n) for assistance in LITERATURE CITED the field. Thanks also to Mario Blanco, Karen Redden, John L. Clark, and H. David Clarke for collecting tissue in BRITTON, N. L. 1915. Studies of West Indian plants. Bulletin of and Guyana. Urs Jauch and Merran Matthews are gratefully the Torrey Botanical Society 42: 372–379. 670 SYSTEMATIC BOTANY [Volume 32

BUREAU, E. 1864. Monographie des Bignoniacees. Paris. 1022 (MO) Ecuador, Gentry, A 14750 (US) Venezuela, Distrito CANDOLLE,A.P.DE. 1838. Revue sommaire de la famille des Federal. Handroanthus chrysotrichus (Mart ex. DC) Mattos Bignoniaceae. Geneve: Bibliotheque Universelle. Keisling 7228 (MO) Argentina, Buenos Aires, Peixoto, A 309 DOS SANTOS, G. and R. B. MILLER. 1992. Wood anatomy of (MO) , Espirito Santo. Handroanthus coralibe (Standl.) Tecomeae. Pp. 336–358 in Flora Neotropica Monograph. 25, Grose Dugand, A 390b (US) Colombia, Atlantico. Han- ed. A. H. Gentry. Bronx: New York Botanical Garden. droanthus guayacan (Seem.) Grose Gentry, A 8574 (US) GENTRY, A. H. 1969. Tabebuia: The tortuous history of a generic , Cayo, Nee, M 10447 (US) , Canal Zone. name (Bignon.). Taxon 18: 635. Handroanthus heptaphyllus (Vell.) Mattos Pedersen, TM ———. 1972. Handroanthus (Bignoniaceae): a critique. Taxon 6526 (US) , Misiones, Gentry, A 51909 (US) Para- 21: 113–114. guay, Paraguari. Handroanthus impetiginosus (Mart ex. DC) ———. 1980. Bignoniaceae- Part I: Crescentieae and Tourret- Mattos Killeen, T 4201 (MO) , Santa Cruz, Nicher 35464 tieae. Flora Neotropica Monograph 25. Bronx: New York (Z) New Caledonia, SEM. Handroanthus obscurus (Bur. & K. Botanical Garden. Schum.) Mattos Schultes, RE 16993 (US) Colombia, Amazonas ———. 1992. Bignoniaceae- Part II (Tribe Tecomeae). Flora Plowman, T 2758 (US) , Loreto. Handroanthus ochraceus Neotropica Monographs 25. Bronx: New York Botanical (Cham.) Mattos subsp. ochraceus Herzog 138 (Z), Bolivia, Garden. SEM. Handroanthus ochraceus (Cham.) Mattos subsp hetero- GROSE,S.O.andR.G.OLMSTEAD. 2007. Evolution of trichus Steyermark, J and Liesner, R 120985 (NY) Venezuela, a charismatic neotropical tree: Molecular phylogeny of Sucre. Handroanthus ochraceus (Cham.) Mattos subsp. Tabebuia s. l. and allied genera (Bignoniaceae). Systematic neochrosanthus Molina, A 1379 (US) , Morazan. Botany 32: 650–659. Handroanthus pulcherrimus (Sandwith) Grose Smith, LB 7263 MATTHEWS, M. L. and P. K. ENDRESS. 2004. Comparative floral (US) Brazil, . Handroanthus pumilus (A. H. structure and systematics in Cucurbitales (Corynocar- Gentry) Grose Hatschenbach, G 49779 (US), Bazil, Han- paceae, Coriariaceae, Tetramelaceae, Datiscaceae, Bego- droanthus serratifolius (Vahl) Grose Broadway, s. n. (Z) niaceae, Cucurbitaceae, Anisophylleaceae). Botanical Grenada, St. Georges, SEM, Zarucchi, JL 4536 (US) Colombia, Journal of the Linnean Society 145: 129–185. Antioquia. Handroanthus subtilis (Sprague & Sandwith) MATTOS, J. R. 1970. Handroanthus, Um novo geˆnero para os Grose Hernandez, L 469 (MO) Venezuela, Bolı´var, A. H. ‘‘ipeˆs’’ do Brasil. Loefgrenia 50: 1–4. Gentry, A 10521 (MO) Venezuela, Bolı´var. Handroanthus MIERS, J. 1863. Report on the plants collected by Mr. Weir, uleanus (Kraenzl.) Grose Stevenson, P 955 (MO). Han- especially the Bignoniaceae. Proceedings of the Royal droanthus umbellatus (Sond.) Mattos A. H. Gentry, A 59133 Horticultural Society 3: 179–202. (NY) Brazil. , Inacio, E 139 (MO) Brazil, MIRANDA, F. 1965. Estudios acerca de arboles y arbustos de Pernambuco. Handroanthus vellosoi (Toledo) Mattos A. H. America Tropical principamente de . Boletin de la Gentry, A 49602 (MO) Brazil, Minas Gerais. Roseodendron Sociedad Botanica de Mexico 29: 34–49. chryseum (Blake) Miranda Aristequieta 6801 (US) Venezuela. RAFFINSQUE, C. 1838. Sylva Telluriana, Philadelphia. Roseodendron donnell-smithii (Rose) Miranda Bullock, SH RHEDER, A. 1913. Neue oder kritische geho¨lze. Mitteilungen 1311 (MO) Mexico, Jalisco, Blanco, C 643 (US) Venezuela, der Deutschen Dendrologischen Gesellschaft 22: 254–265. Bolivar, S. Grose 165 (WTU), Me´xico, Chiapas, SEM. Tabebuia ROSE, J. N. 1892. A new species from Mexico and Central aurea (Manso) Benth and Hook. f.ex. S Moore Foster, P 163 America Tabebuia donnell smithii sp.nov. Botanical Gazette (MO) Bolivia, Santa Cruz, Thomas, W et al 4646 (US) Brazil, 17: 418, Pl. 426. Matto Grosso. Tabebuia cristata A. H. Gentry Peixoto, A 3067 SCHUMANN, K. 1894. Bignoniaceae. Pp. 189–252 in Die (US) Brazil, Espirito Santo. Tabebuia insignis (Miq.) Sand- Natu¨rlichen Pflanzenfamilien, Vol 4, eds. A. Engler and with var. orinocensis Maguire, B. 37701 (US) Venezuela, K. Prantl. Leipzig. Amazonas. Tabebuia insignis (Miq.) Sandwith var. insignis SEEMAN, B. 1863. Revision of the natural Bignoniaceae. Henkel, T 723 (US) Guyana, Potaro-Siparuni. Tabebuia Seemann’s Journal of Botany 1: 18–19. insignis (Miq) Sandw. var. monophylla Gillespie, L. J. 927 SEIBERT, R. J. 1940. The Bignoniaceae of the Maya area (US) Guyana, Potaro-Siparuni. Tabebuia microphylla (Lam.) including Yucatan, Campeche, Quintana Roo, Chiapas, Urb. Zanoni, T 20808 (NY) Dominican Republic, Pedernales, Tabasco, British Honduras and . Carnegie Zanoni, T 36525 (MO) Dominican Republic, Pedernales. Institute of Washington Publications 522: 357–434, pls. Tabebuia myrtifolia (Greenm.) A. H. Gentry var. petrophila 355–357. Leon 11489 (NY) Cuba, Havana. Tabebuia myrtifolia SPANGLER, R. E. and R. G. OLMSTEAD. 1999. Phylogenetic (Greenm.) A. H. Gentry var. myrtifolia Garcia, R 510 (MO) analysis of Bignoniaceae based on the cpDNA gene Dominican Republic, Pedernales. Tabebuia nodosa (Griseb.) sequences rbcL and ndhF. Annals of the Missouri Botanical Griseb. Hatschenbach G 49232 (US) Brazil, Curitiba. Tabebuia Garden 86: 33–46. ophiolitica Alain A. H. Gentry A 50633 (MO) Republica SPRAGUE, T. A. and N. Y. SANDWITH. 1932. of British Dominica, Puerto Plata. Tabebuia orinocensis (Sandwith) A. Guiana and Trinidad. Kew Bulletin 1932: 18–29. H. Gentry Gentry, A. 14609 (US) Venezuela, Amazonas, Davidse, G. 15160 (US) Venezuela, Atures. APPENDIX 1. List of representative herbarium specimens (Lindl.) Miers Nicholson, DH 1987 (US) Dominica, Wilbur, RL consulted for mophology. 7518 (US) Dominica. Tabebuia palustris Hemsl. A. H. Gentry, Handroanthus albus (Cham.) Mattos Reitz and Klein 7037 A 7376 (MO) Colombia, Choco´. Tabebuia pilosa A. H. Gentry (US) Brazil, Santa Catarina, Hatschbach, G 48929 (US) Brazil, Cuello, N 723 (US) Venezuela, Bolivar, Wurdack, JJ 39990 (US) Parana. Handroanthus barbatus (E.Mey) Mattos Prance, GT Venezuela, Bolivar. Tabebuia rigida Urban Axelrod, F 4107 20577 (US) Brazil, Amazonas, Maguire and Wurdack 34873 (US) Puerto Rico, Puerto Rico. Tabebuia rosea (Bertol.)DC (US) Venezuela, Amazonas. Handroanthus billbergii (Bur & Morton, CV 2691 (US) Me´xico, Oaxaca, Breteler, FJ 4418 (US) K. Schum) Grose subsp ampla Steyermark, J. 94543 (US) Venezuela, Zu´lia, S. Grose 156 (WTU), Me´xico, Veracruz, Venezuela, Falcon Samaniego, V 015 (US) Ecuador Loja. SEM. Tabebuia roseo-alba (Ridley) Sandwith Prance, GT, Handroanthus capitatus (Bur. & K. Schum) Mattos Balee, W Lleras,E, and Coelho, DF 19034 (NY) Brazil, Mato Grosso. 5097 (MO) Bolivia, Beni, Blanco, C 496 (US) Venezuela, Delta Sprague & Stapf de Bruijn 1633 (US) Amacuro. Handroanthus chrysanthus (Jacq.) Grose Gentry, A Venezuela, Bolı´var.