Systematic Botany (2007), 32(3): pp.

660–670
# Copyright 2007 by the American Society of Plant Taxonomists

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.
1
Author for correspondence (sheydler@smith.alumnae.net)

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 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 the
leaves and calyx. The species of Handroanthus are also characterized by having extremely dense wood containing
copious quantities of lapachol. Tabebuia is restricted to those species with white to red or rarely yellow flowers 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, taxonomy.

Tabebuia, as currently circumscribed with 100
species, is the largest genus in Bignoniaceae and is
distributed from the southwestern U.S. to northern
Argentina and Chile. Over the course of its
taxonomic history, it has been split and re-
assembled several times, as researchers interpreted
the morphological diversity in different ways. The
wide range of morphological diversity suggests
there may be more than one lineage included
within the traditional concept of Tabebuia. Recent
studies (Spangler and Olmstead 1999; Grose and
Olmstead 2007) have confirmed that Tabebuia, as
currently delimited, is polyphyletic. Therefore, the
generic boundaries need to be redefined in
Tabebuia and related genera. The goal of this paper
is to revise the generic classification to be consis-
tent with its phylogeny.
The name Tabebuia has a long and convoluted
history (Gentry 1969). This name was first published
by de Candolle (Candolle 1838) who applied it to
bignoniaceous trees with ‘‘simple’’ leaves. Howev-
er, the concept of Tabebuia came to be expanded to
encompass a large amount of morphological di-
versity. As researchers examined and monographed
this taxon, a number of concepts of Tabebuia
emerged, producing a labyrinthine synomomy
(Candolle 1838; Raffinsque 1838; Sprague and
Sandwith 1932). In addition, there was confusion
as to the boundaries between Tecoma and Tabebuia
(Miers 1863; Seeman 1863; Bureau 1864; Schumann
1894; Rheder 1913). Much of the confusion between
Tecoma and Tabebuia was sorted out by Britton
(Britton 1915). However, disagreement as to the
660
delimitation of Tabebuia has continued into recent
times (Mattos 1970; Gentry 1972).
Mattos (1970) divided Tabebuia into two different
groups, indicating that a group of Brazilian taxa
known as ‘‘îpes’’ should not remain in Tabebuia.
His reasoning appears to have been based on the
original concept for Tabebuia, which he defined as
having simple leaves and an ovary that in cross
section has 3–4 ovules/locule. This morphology is
consistent with the type of Tabebuia, T. cassinoides,
but is the most exclusive concept for Tabebuia since
it was proposed by de Candolle (1838). Neither did
Mattos think the îpes belonged in Tecoma, due to
their palmately compound leaves. He created the
genus Handroanthus for those species with pal-
mately compound leaves and 8–9 series of ovules/
locule, and typified this genus with Handroanthus
albus (Tabebuia alba).
Gentry (1972, 1992) included Handroanthus in
Tabebuia, insisting that Tabebuia as previously de-
lineated is natural. Gentry (1972) was adamant that
the large genus Tabebuia should not be broken up
further, and expressed his ‘‘sincere hope that future
students of Bignoniaceae will consider the case of
Handroanthus before succumbing to further parox-
ysms of unwarranted splitting’’ (Gentry 1972).
Gentry (1992) recognized some groups within
Tabebuia, however. In his treatment for Flora
Neotropica he defined 10 species groups although
he placed two species, T. arimaoensis and T.
heterophylla, in both groups 9 and 10. The largest
of Gentry’s groups are primarily species endemic
to the Greater Antilles (55 species). He did not use
2007] GROSE & OLMSTEAD: REVISION OF TABEBUIA
phylogenetic terminology to describe the relation-
ships, but he clearly believed that this group of
Anillean species was monophyletic and sister to
a group of mainland species that are spread
throughout the continental part of the genus’ range
(Gentry 1992).
Tabebuia donnell-smithii was placed in Tabebuia by
Rose (1892) with some reservation: ‘‘The species,
while not agreeing in all respects with Tabebuia,
answers better to this than to any other known
genus. In its inflorescence and ribbed pods it is
more like Godmania or Cybistax but does not agree
in other particulars.’’ This species was later moved
to Cybistax by Seibert (1940). However, Miranda
(1965) transferred this species to a new genus,
Roseodendron, segregating it from Cybistax on the
basis of calyx texture and shape, sessile nature of
its 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 all
Tabebuia species and had representatives from all
10 of Gentry’s (1992) species groups. The findings
showed that Tabebuia consists of three clades.
‘‘Tabebuia goup I’’ is sister to the Caribbean
endemic Ekmanianthe. ‘‘Tabebuia group II’’ is sister
to Crescentieae+Spirotecoma. The third clade is
represented by T. donnell-smithii and is sister to
Tabebuia group II +Crescentieae+ Spirotecoma.
MATERIALS AND METHODS
Characters to support the phylogeny of Grose and
Olmstead (2007) are based on herbarium holdings at US,
MO and NY and published morphological and anatomical
studies (Gentry 1992; dos Santos and Miller 1992). Herbarium
specimens were consulted for morphological characters
(Appendix 1). Leaf surfaces and calices were examined with
a scanning electron microscope (SEM). Specimens were
prepared for the SEM according to the protocol established
in Matthews and Endress (2004)
MORPHOLOGICAL CHARACTERS DEFINING CLADES
Each of the three clades of Tabebuia: Tabebuia
groups I and II and T. donnell-smithii is supported
by several morphological characters. Gentry de-
scribed the importance of the calyx in Bignoniaceae
taxonomy especially in tribe Bignonieae (Gentry
1980). This study confirms previous observations
that calyx morphology as well as indumentum,
wood anatomy, flower color, and fruit morphology
have been important in defining the three groups
of Tabebuia in this clade (dos Santos and Miller
1992).
Tabebuia group I shares the characters of an
indumentum of lepidote scales and a spathaceous,
sometimes irregularly bi- or trilabiate calyx
661
(Fig. 1C, F). The lepidote scales in a group of
species (such as Tabebuia pilosa) may be stalked.
The wood anatomy of Tabebuia group I is distinc-
tive; it is lightweight, with medium specific gravity
of 0.4–0.74, and lacks lapachol (dos Santos and
Miller 1992). The heartwood is indistinct from the
sapwood and the rays are 1–2 cells in width (dos
Santos 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 (Gentry
1992). The flowers of Tabebuia group I are white to
red in color, and often have a yellow throat (Fig 1I).
Two species belonging to this clade, Tabebuia aurea
and the unsampled Tabebuia nodosa have yellow
flowers but otherwise share the above character-
istics.
Tabebuia group II can be distinguished by having
an indumentum of simple to stellate or dendroid
hairs (Fig. 1A, D). Lepidote scales are also usually
present, but may be obscured by the hairs. The
calyx is campanulate to cupular and 5-dentate
(Fig. 1D). The wood is among the heaviest and
hardest known, with a basic specific gravity
greater than 0.74 (dos Santos and Miller 1992). It
is also distinctive among Tabebuia s. l., in having
very large pits (ranging from 7–14 mm) in their
vessel elements (dos Santos and Miller 1992). Only
two species in this group, T. pumila and T.
selachidentata, are described as occasionally having
unifoliolate leaves (Gentry 1992). In all other cases,
the leaves are digitately 3–9 foliolate. The fruits in
this group are linear and slightly costate to smooth,
and often are densely tomentose. The flowers in
Tabebuia group II are generally yellow (Fig. 1G) or,
in four species, T. selachidentata, T. barbata, T.
heptaphylla and T. impetiginosa, magenta with
yellow throat (Gentry 1992).
Tabebuia donnell-smithii represents a group dis-
tinguished by its unique calyx (Fig. 1E). It is
spathaceous in shape, and of the same color and
texture as the corolla. This lineage contains only
one other species, T. chrysea. The indumentum is of
lepidote scales and small, glandular hairs (Fig. 1B).
The wood is similar to that of Tabebuia group I but
contains tyloses, or intrusive growths of the cell
wall into the vessel cavities, and pits intermediate
in 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, and
fruits are linear and irregularly costate and bullate.
TAXONOMIC CONSEQUENCES OF THE PHYLOGENY
The three clades of Tabebuia s. l. correspond
closely to the aggregations of species groups of
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 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 showing
lepidote scales. Scale bars for each photograph as indicated. D–F. Calyx shapes and flower color. D. Handroanthus chrysanthus
2007] GROSE & OLMSTEAD: REVISION OF TABEBUIA 663

recognized as genera here. The type of Tabebuia, T.
cassinoides (Lamarck) A.P. de Candolle, was not
included in the molecular study due to the
unavailability of fresh material and the lack of
success in extracting DNA from herbarium speci-
mens. However, it has simple leaves and an
indumentum of lepidote scales covering the leaves
and calyx, which places it confidently in Tabebuia
group I. We now restrict Tabebuia to those species
occurring in this clade. This corresponds to
Gentry’s (1992) informal groups 2 and 6–10 of
Tabebuia. The sister taxon to this clade, Ekma-
nianthe, shares the characters described for Tabebuia
group I. It has 9-foliolate leaves, lepidote scales,
and has similar wood anatomy; nevertheless, it is
morphologically distinct by virtue of its hawk-
moth-pollinated flowers having long corolla tubes
with united lobes, and basally curved fruit. The
autapomorphy for Tabebuia I exclusive of Ekma-
nianthe is its straight, cylindrical fruit.
The second clade, Tabebuia group II, comprises
most of the species with yellow-flowers, and
includes 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 study
due to lack of available tissue, but it shares the four
characters listed above and is assignable unambig-
uously to this clade. Gentry (1972) insisted that
Couralia 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 to
Tabebuia I, so the name Couralia remains a synonym
for Tabebuia. Accordingly, priority rests with
Handroanthus 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 resurrected
for T. donnell-smithii and T. chrysea, since the latter,
like T. donnell-smithii has calyces that are texturally
identical to the corolla. These two species form
Gentry’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, lapachol
present 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 at
least along vein axils and sometimes densely
covering leaves; petiolule to 9 cm; petiole to
30 cm. Inflorescence dichotomously branched,
without a well developed central rachis, sometimes
contracted; 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 tomentose
with 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 mm
in H. subtilis) staminode reduced. Ovary conical to
linear-oblong; ovules 2–10 seriate in each locule.
Fruit an elongate linear to cylindric capsule,
smooth to slightly costate, glabrous to scattered
lepidote to pubescent; pubescence scattered to
villous; trichomes simple, stellate or dendroid.
Seeds thin, bialate; wings hyaline membranacous
and sharply demarcated from seed body.
A genus of 30 species distributed throughout
Central and South America with one species (H.
billbergii) in the Antilles.
Taxa transferred to Handroanthus (*indicates the
nine 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. Paraná 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 Neotropica
25(2): 142. 1992.

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
3. H. BARBATUS (E. Mey.) Mattos, Loefgrenia 50: 2.
1970. Bignonia barbata E. Mey. Nova Acta
Phys.-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. Tabebuia
barbata (E. Meyer) Sandwith, Lilloa 3: 462.
1938.
Bignonia fluviatilis Aublet sensu Humboldt, Bon-
pland and Kunth, Nov. gen. sp. Pl3: 139. 1819
non Aublet.
Tecoma toxophora Martius emend DC, Prodr. 9: 217.
1845 (excl. sin. Margr.) non Martius, Flora 24,
Beibl. 15. 1841. Couralia toxophora (Martius
emend 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. in
Martius, Fl. Bras. 8(2): 319. 1897.
a. H. billbergii. subsp. billbergii. Tabebuia bill-
bergii subsp. billbergii Bureau & K. Schum. in
Martius, 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, Flora
Neotropica 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 Tabebuia
dentata, 1863. Handroanthus dentatus (Bur. and
K. 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. miquelii
DC, Prodr. 9: 219. 1845.
Tabebuia glomerata Urban, Feddes Rebert. 14: 305.
1916.
Tabebuia hypolepra Sprague and Sandwith, Kew
Bull. 1932: 25 1932.
8. H. catarinensis (Gentry) S. Grose comb. nov.
Tabebuia catarinensis Gentry, Ann. Missouri
Bot. 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.
[Volume 32
a. H. chrysanthus subsp. chrysanthus*
Tabebuia rufescens J. R. Johnston, Proc. Amer.
Acad. Arts 40: 696. 1905.
Tecoma evenia Donn. Sm., Bot. Gaz. (Crawfords
ville) 20. 1895 pro parte.
Tecoma palmeri Kraenzl. Feddes Repert, 17: 220.
1921.
b. H. chrysanthus subsp. meridionalis (A. H.
Gentry) S. Grose, comb. nov. Tabebuia
chrysantha ssp. meridionalis A. Gentry, Phy-
tologia 35: 193. 1977.
Tecoma spectabilis Planchon and Linden Fl. Serres
Jard. L’Europe. 9: 233. 1854.
Tabebuia spectabilis (Planch. and Linden) Ni
chols., Ill. dict. gard. 4: 1. 1887.
c. H. chrysanthus subsp. pluvicolus (A. H.
Gentry) S. Grose comb. nov.* Tabebuia
chrysantha (Jacquin) Nicholson subsp. pluvi-
cola. A. Gentry in Phytologia 35: 190. 1977.
Tecoma grandis Appun., Behand. Samereien und
Pflanzen 39. 1858, nom. nud.
10. H. CHRYSOTRICHUS (Mart. ex DC) Mattos,
Loefgrenia 50: 2. 1970.* Tecoma chrysotricha
Mart. ex. DC, Prodr. 9: 216. 1845. Gelseminum
chrysotrichum (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 São 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 Neotropica
25(2): 174. 1992.
13. H. guayacan (Seeman) S. Grose comb. nov.*
Tecoma guayacan Seemann, Bot. voy. Herald
180. 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.
2007] GROSE & OLMSTEAD: REVISION OF TABEBUIA
251. 1829. Tabebuia heptaphylla (Vell.) Toledo,
Arq. Bot. Estado São Paulo, n. s. 3: 33. 1952.
Tecoma eximia Miq., Linnaea 22: 803. 1849. Tabebuia
eximia (Miq.) Sandwith, Lloydia 2: 213. 1939.
Handroanthus eximia (Miq.) Mattos, Loefgrenia
50: 2. 1970.
Tecoma curialis Saldanha da Gama, Config, descr.
orgãos 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 São Paulo, n. s., 3: 1952.
Tabebuia impetiginosa var. lepidota (Bur.) Toledo,
Arq. Bot. Estado São 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 ex
Griseback) 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 impetiginosus
var. 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. Genève 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.
665
Tabebuia schunkevigoi Simpson, Fieldiana, Bot. 36: 1.
1972.
16. H. incanus (A. H. Gentry) S. Grose comb. nov.
Tabebuia incana A. Gentry in Ann. Missouri
Bot. 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. Venezuela
8(4): 391. 1982.
b. H. OCHRACEUS subsp. NEOCHRYSANTHUS (A. H.
Gentry) S. Grose.* Tabebuia neochrysantha A.
H. Gentry, Brittonia 22: 260. 1970. Tabebuia
ochracea (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. Tabebuia
hypodiction (DC) Standl., Field Mus. Nat. Hist.,
Bot. Ser. 11: 176. 1936.
Tecoma heteropoda DC, Prodr. 9: 219. 1845. Tabebuia
ochracea subsp. heteropoda (DC) A. H. Gentry,
in Prance, Biological Diversification in the
Tropics 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 Kränzl., Feddes Repert. 17: 216. 1921.
Tecoma campinae Kraenzlin, Feddes Repert. 17: 215.
1921.
Tecoma hemmendorffiana Kraenzl., Feddes Repert.
17: 224. 1921.
666 SYSTEMATIC BOTANY
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 Neotropica
25(2): 244. 1992.
23. H. riodocensis (A. H. Gentry) S. Grose comb.
nov. Tabebuia riodocensis A. H. Gentry, Flora
Neotropica 25(2): 1248. 1992.
24. H. selachidentatus (A. H. Gentry) S. Grose
comb. 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 and
Britton, 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. Bignonia
conspicua Richard ex DC, Prodr. 9: 221, nom.
nud., pro syn.
Tecoma patrisiana DC, Prodr. 9: 221. 1845.
Tecoma speciosa DC, Prodr. 9: 218. 1845. Gelseminum
speciosum (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
[Volume 32
comb. nov.* Tabebuia subtilis Sprague and
Sandwith, Kew Bull. 1932: 23. 1932.
28. H. uleanus (Kraenzl) S. Grose comb. nov.
Tecoma uleana Kränzlin, 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. Tabebuia
eximia (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 São 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. Estado
São Paulo, n. s. 3(1): 34. 1952.
Bignonia longiflora Vell., Fl. flumin. 252. 1829 (1825).
Tecoma longiflora (Vell.) Bur. & K. Schum., in
Martius, Fl. Bras. 8(2): 324. 1897, non Tecoma
longiflora Griseb. 1866.
Tecoma alba Cham. var. subdenudata Bur., Dansk.
Naturhist. Foren. Kjøbenhavn 1893: 115. 1893.
ROSEODENDRON Miranda, Bol. Soc. Not. Mex. 29:43
1965.—TYPE: Roseodendron donnell-smithii
(Rose) Miranda.
Trees to 35 m; heartwood not very distinct from
sapwood, yellowish or light to reddish brown;
tyloses present; lapachol present in small quanti-
ties (R. chrysea); rays 2–5(8) cells wide; medium
intervessel pits (6–8 mm); thin to thick walled
fibers; medium specific gravity (0.4–7.4). Leaves
palmately 5–7 foliolate; leaflets oblong-elliptic, to
28 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 dendroid
trichomes, or more or less glabrate, drying olive,
darker above and lighter below; apex acute to
acuminate; bases truncate; petiolules to 7 cm long;
petiole to 26 cm long, tomentose with dendroid
and barbate trichomes or glabrate. Inflorescence
a large terminal panicle with the central rachis well
developed and lateral branches either very short
and dendroid-pubescent, or branching with rachis
and branches puberulous with capitate trichomes.
Flowers: Calyx campanulate, 10–20 mm long, 5–
15 mm wide, finely membranaceous, bilabiate to
irregularly several–5-dentate, somewhat lepidote
2007] GROSE & OLMSTEAD: REVISION OF TABEBUIA
and puberulous, sometimes with gland tipped
trichomes. Corolla yellow, sometimes with reddish
lines in throat, tubular-infundibuliform; tube 3–
4.5 cm long, 0.8–2 cm wide at mouth of tube, lightly
puberulous to almost glabrous outside, generally
glabrous inside, pubescent at level of stamen
insertion; lobes glabrous to lightly pubescent; lobes
1–2.5 cm long. Stamens didynamous; thecae di-
varicate, 2–2.5 mm long; staminode reduced. Ovary
linear, 5–6 mm long, 0.6–1.5 mm wide; ovules 4 or 6
seriate in each locule. Fruit an elongate-linear
capsule, 25–45 cm long, 0.9–3 cm wide, longitudi-
nally striate-costate to irregularly striate costate with
8–12 rather irregular longitudinal ribs, puberulous
with simple and branched or simple-gland-tipped
trichomes. Seeds thin; wings hyaline-membranac-
eous and sharply demarcated from seed body.
Two species ranging from México to Northwest-
ern South America, and dry areas of Venezuela
and Colombia.
Taxa transferred to Roseodendron (* indicates
taxon 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 Woods
63: 7. 1940.
2. R. DONNELL-SMITHII (Rose) Miranda, Bol. Soc.
Mex. 29: 43. 1965.* Tabebuia donnell-smithii
Rose, Bot. Gaz. 17: 418. pl. 26. 1892. Cybistax
donnell-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 Genève, ser 2, 17:130. Sep. 1838.—
TYPE: Tabebuia uliginosa (Gomes) DC 5
Tabebuia cassinoides (Lam.) DC, Encyc. Méth.
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 obtusifolia
Lam.) 5 T. cassinoides (Lamarck) DC
Couralia Splitgerberger, Tujdschr. Natuurl. Gesch.
Physiol. 9: 14. 1842.—TYPE: Couralia fluviatilis
(Aubl.) Splitgerberger 5 T. fluviatilis (Aubl.) DC.
667
Trees or shrubs; heartwood not distinct from
sapwood, light to reddish brown; lapachol absent;
rays 1–3 cells wide; small to medium intervessel
pits (3–6 mm); thin to thick walled fibers; medium
specific gravity (0.4–7.4). Leaves 1–7(9) foliolate;
leaflets narrowly elliptic to orbicular to 35 cm long
and 32 cm wide, with stalked or sessile lepidote
scales that are usually scattered and sometimes
densely covering undersurface of leaves; petiolule
to 11 cm; petiole to 18 cm. Inflorescence dichoto-
mously branching, without a well developed central
rachis, usually a few flowered panicle, occasionally
many 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 of
stamen insertion. Stamens didynamous; thecae
divaricate, 2–6 mm; staminode reduced. Ovary
linear; ovules 2–3 seriate in each locule. Fruit an
elongate linear to cylindric capsule, smooth to
costate striate, minutely to densely lepidote. Seeds
thin, bialate; wings hyaline membranacous and
sharply demarcated from seed body.
A genus of 67 species widely distributed
throughout Central and South America and the
Antilles.
Included taxa (* indicates taxon used in Grose
and Olmstead 2007). Names in this group remain
unchanged from Gentry (1992) and are listed for
convenience:
1. TABEBUIA ACROPHYLLA (Urb.) Britton*
2. T. ANGUSTATA Britton
3. T. ARIMAOENSIS Britton
4. T. AUREA (Silva Manso) Benth. & Hook. f. ex S.
Moore*
5. T. BAHAMENSIS (Northr.) Britton*
6. T. BERTEROI (DC) Britton*
7. T. BIBRACTEOLATA (Griseb.) Britton
8. T. BROOKSIANA Britton
9. T. BUCHII (Urb.) Britton
10. T. BULLATA A. H. Gentry
11. T. CALCICOLA Britton
12. T. CALETICANA A. H. Gentry and D. Albert
13. T. CASSINOIDES (Lam.) DC (Type)
14. T. CLEMATIS Alain
15. T. CONFERTA Urb.
16. T. CRISPIFLORA Alain
17. T. XDEL-RISCOI Borhidi
18. T. DENSIFOLIA Urb.
19. T. DOMINGUENSIS (Urb.) Britton
20. T. DUBIA (C.Wright ex Sauvalle) Britton ex
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. (Miq.) Sandwith var. INSIGNIS*
INSIGNIS 60. T. SCHUMANNIANA Urb.*
b. T. INSIGNIS(Miq.) Sandwith var. MONO- 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 Key to the Genera. 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 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sparattosperma
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 raceme, 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 forests 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. Plants shrubs or subshrubs, 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 Cerrado . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 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. Inflorescences cauliflorous or ramiflorous, growing from round or branchlike areas on bark (terminal in some
species of Amphitecna) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 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 pedicel elongate; corolla yellow to
dark red, lower corolla lobes reflexed; stamens exserted, 4 spp. Cuba and Hispañola . . . . . . . . . . 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. México 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, tearing
irregularly. 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, then
mangrove 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 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 Hispañola. . . . 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
dissertation of the first author. Thanks to Michelle Zjhra, Lúcia
Lohmann, and one anonymous reviewer for comments that
significantly improved this manuscript and to Daisy Castillo
(JBSD), Theodoro Clase (JBSD), Alberto Veloz (JBSD), Frank
Axelrod (UPRRP), Javier Francisco-Ortega (FBG/FIU), José
Gonzalez (INBio), Mónica Mejı́a-Chang, Carmen Galdames
(STRI), Carlos Burelo Ramos (XAL), Ricardo Rueda and Dania
Paguaga (Universidad de Nicaragua, León) for assistance in
the field. Thanks also to Mario Blanco, Karen Redden, John L.
Clark, and H. David Clarke for collecting tissue in Ecuador
and Guyana. Urs Jauch and Merran Matthews are gratefully
acknowledged for their support with the SEM. NYBG, MO
and US kindly loaned specimens for this study. Funds for this
research were provided by NSF DEB: 0309065, OTS, the
Garden Club of America, ASPT, BSA, University of Washing-
ton, Department of Botany ‘‘Plant Molecular Systematics
Fellowship’’, and the University of Washington Department
of Biology ‘‘Melinda Denton Fellowship’’.
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APPENDIX 1. List of representative herbarium specimens
consulted 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, GT
20577 (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, W
5097 (MO) Bolivia, Beni, Blanco, C 496 (US) Venezuela, Delta
Amacuro. Handroanthus chrysanthus (Jacq.) Grose Gentry, A
[Volume 32
1022 (MO) Ecuador, Gentry, A 14750 (US) Venezuela, Distrito
Federal. Handroanthus chrysotrichus (Mart ex. DC) Mattos
Keisling 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, TM
6526 (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, Amazonas
Plowman, 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, Bolı́var, A. H.
Gentry, A 10521 (MO) Venezuela, Bolı́var. Handroanthus
uleanus (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. Roseodendron
chryseum (Blake) Miranda Aristequieta 6801 (US) Venezuela.
Roseodendron donnell-smithii (Rose) Miranda Bullock, SH
1311 (MO) Mexico, Jalisco, Blanco, C 643 (US) Venezuela,
Bolivar, S. Grose 165 (WTU), México, Chiapas, SEM. Tabebuia
aurea (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. insignis
Henkel, T 723 (US) Guyana, Potaro-Siparuni. Tabebuia
insignis (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. petrophila
Leon 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. Tabebuia
ophiolitica Alain A. H. Gentry A 50633 (MO) Republica
Dominica, 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, RL
7518 (US) Dominica. Tabebuia palustris Hemsl. A. H. Gentry,
A 7376 (MO) Colombia, Chocó. Tabebuia pilosa A. H. Gentry
Cuello, 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.)DC
Morton, CV 2691 (US) México, Oaxaca, Breteler, FJ 4418 (US)
Venezuela, Zúlia, S. Grose 156 (WTU), México, 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, Bolı́var.