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strong>Taxonomic Revision of Tynanthus (Bignonieae, Bignoniaceae)</strong

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Tynanthus is a genus of Neotropical lianas that are distributed from southern Mexico to southern Brazil. Extensive herbarium work, field studies and new molecular phylogenetic data were used as the basis to recognize 14 species in the genus. Here, we provide a detailed treatment of the group, including an identification key to all species recognized, as well as morphological descriptions, a complete list of synonymy, nomenclatural information, taxonomic notes, phenological data, information on habitat and distribution, and illustrations. Two species are circumscribed differently from earlier classifications, in particular Tynanthus elegans is synonymized with Tynanthus cognatus, while Tynanthus villosus is synonymized with Tynanthus polyanthus. In addition, Tynanthus goudotianus is treated as a doubtful name, and two recently described species are recognized, Tynanthus densiflorus and Tynanthus espiritosantensis. One species name is neotypified, Tynanthus caryophylleus, and nine species names are lectotypified, namely Cuspidaria ovalis, Tynanthus cognatus, Tynanthus elegans, Tynanthus guatemalensis, Tynanthus lindmanii, Tynanthus myrianthus, Tynanthus panurensis, Schizopsis chimonantha, and Schizopsis regnelliana. A complete list of doubtful and excluded names is presented.
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Accepted by Peter Heenan: 23 Feb. 2015; published: 19 Jun. 2015
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PHYTOTAXA
ISSN 1179-3155 (print edition)
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1179-3163
(online edition)
Copyright © 2015 Magnolia Press
Phytotaxa 216 (1): 001–060
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phytotaxa
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http://dx.doi.org/10.11646/phytotaxa.216.1.1
PHYTOTAXA
Taxonomic Revision of Tynanthus (Bignonieae, Bignoniaceae)
MARIA CLÁUDIA MELO PACHECO DE MEDEIROS
1,2
& LÚCIA G. LOHMANN
1,2
1 Universidade de São Paulo, Instituto de Biociências, Departamento de Botânica, Rua do Matão, 277, 05508–090, São Paulo, SP,
Brazil.
2 Authors for correspondence: mariaclaudiamedeiros@hotmail.com; llohmann@usp.br
Magnolia Press
Auckland, New Zealand
216
MEDEIROS & LOHMANN
2
Phytotaxa 216 (1) © 2015 Magnolia Press
MARIA CLÁUDIA MELO PACHECO DE MEDEIROS & LÚCIA G. LOHMANN
Taxonomic Revision of Tynanthus (Bignonieae, Bignoniaceae)
(Phytotaxa 216)
60 pp.; 30 cm.
19 June 2015
ISBN 978-1-77557-727-0 (paperback)
ISBN 978-1-77557-728-7 (Online edition)
F
IRST
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UBLISHED I
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2015 B
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ISSN 1179-3155 (Print edition)
ISSN 1179-3163 (Online edition)
Phytotaxa 216 (1) © 2015 Magnolia Press
3
TAXONOMIC REVISION OF TYNANTHUS (BIGNONIEAE)
Abstract
Tynanthus is a genus of Neotropical lianas that are distributed from southern Mexico to southern Brazil. Extensive
herbarium work, field studies and new molecular phylogenetic data were used as the basis to recognize 14 species in the
genus. Here, we provide a detailed treatment of the group, including an identification key to all species recognized, as
well as morphological descriptions, a complete list of synonymy, nomenclatural information, taxonomic notes,
phenological data, information on habitat and distribution, and illustrations. Two species are circumscribed differently
from earlier classifications, in particular Tynanthus elegans is synonymized with Tynanthus cognatus, while Tynanthus
villosus is synonymized with Tynanthus polyanthus. In addition, Tynanthus goudotianus is treated as a doubtful name,
and two recently described species are recognized, Tynanthus densiflorus and Tynanthus espiritosantensis. One species
name is neotypified, Tynanthus caryophylleus, and nine species names are lectotypified, namely Cuspidaria ovalis,
Tyn anthu s cog n atus, Tynant h us e legan s , Tynanthus guatemalens i s , Ty n anthu s l i ndman ii, Tyn a nthus myria n thus,
Tynanthus panurensis, Schizopsis chimonantha, and Schizopsis regnelliana. A complete list of doubtful and excluded
names is presented.
Resumo
Tynanthus é um gênero de lianas Neotropicais distribuídas desde o sul do México até o sul do Brasil. Trabalho extenso
em herbário, estudos de campo e novos dados moleculares foram utilizados como base para reconhecer 14 espécies no
gênero. Aqui, nós fornecemos um tratamento detalhado do grupo, incluindo uma chave de identificação de todas as
espécies reconhecidas, bem como descrições morfológicas, uma lista completa de sinonímias, informações
nomenclaturais, comentários taxonômicos, dados fenológicos, informações sobre habitat e distribuição e ilustrações.
Duas espécies são circunscritas diferentemente de classificações anteriores, em particular Tynanthus elegans é
sinonimizado em Tynanthus cognatus, enquanto Tynanthus villosus é sinonimizado em Tynanthus polyanthus.
Adicionalmente, Tynanthus goudotianus é tratado como nome duvidoso, e duas espécies descritas recentemente são
reconhecidas, Tynant hus densi fl oru s e Tynanthus espiritosantensis. Um nome de espécie é neotipificado, Tynanthus
caryophylleus, e nove nomes de espécies são lectotipificados, a saber: Cuspidaria ovalis, Tynanthus cognatus, Tynanthus
elegans, Tyn ant hus guat emalen sis, Tynanthu s lindma nii , Tynanthu s myriant hus, Tyn ant hus panurensis, Schizopsis
chimonantha e Schizopsis regnelliana. Uma lista completa de nomes duvidosos e excluídos é apresentada.
Key words: “cipó-cravo”, “clavo huasca”, Neotropical flora
Table of contents
Introduction .................................................................................................................................................................................. 4
Material and methods .................................................................................................................................................................... 5
Results ........................................................................................................................................................................................... 5
Taxonomic treatment ..................................................................................................................................................................... 5
Tyn an thus .............................................................................................................................................................................. 5
Key to species of Tyna nth us .................................................................................................................................................. 6
1. Tynanthus cognatus............................................................................................................................................................ 7
2. Tynanthus croatianus ...................................................................................................................................................... 12
3. Tynanthus densiflorus ...................................................................................................................................................... 14
4. Tynanthus espiritosantensis ............................................................................................................................................. 16
5. Tynanthus fasciculatus .................................................................................................................................................... 19
6. Tynanthus guatemalensis ................................................................................................................................................. 22
7. Tynanthus labiatus ........................................................................................................................................................... 25
8. Tynanthus macranthus ..................................................................................................................................................... 28
9. Tynanthus micranthus ..................................................................................................................................................... 30
10. Tynanthus panurensis .................................................................................................................................................... 33
11. Tynanthus polyanthus .................................................................................................................................................... 35
12. Tynanthus pubescens ..................................................................................................................................................... 43
13. Tynanthus sastrei ........................................................................................................................................................... 44
14. Tynanthus schumaniannus ............................................................................................................................................. 47
Doubtful and excluded names ............................................................................................................................................. 51
MEDEIROS & LOHMANN
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Phytotaxa 216 (1) © 2015 Magnolia Press
Acknowledgements ..................................................................................................................................................................... 51
References ................................................................................................................................................................................... 51
Appendix. Index to specimens examined ................................................................................................................................... 53
Introduction
Tyn anthus Miers (1863: 193) includes 14 species, and consists on a small genus of Neotropical lianas. The genus is
distributed from southern Mexico to southern Brazil, where it predominates in humid forests, although few
representatives are also found in dry areas. Species of Tynanthus generally present restricted geographic
distribution, occurring in Western South America, Central America, Amazonia or Atlantic Forest exclusively
(Lohmann & Taylor 2014).
The genus was described ca. 150 years ago by Miers (1863), who originally included nine species in the
genus. From those, only four names were validly published, T. co gnatus (Chamisso 1832: 703) Miers (1863: 193),
T. elegans Miers (1863: 193), T. fasciculatus (Vellozo 1825: 247; 1827: tab. 25) Miers (1863: 193), and T. labiatus
(Chamisso 1832: 701) Miers (1863: 193). One year after the description of the genus, Bureau (1864), without
noticing the earlier publication of Tynanthus, described the new genus Schizopsis Bureau (1864: 44) in which he
subsequently included seven species (Bureau 1865). Three years later, Bureau (1868) had already noticed the
overlap between Tynanthus and Schizopsis, and published a taxonomic revision of Tynanthus in which he proposed
the appropriate synonymizations and combinations [e.g., T. gou dotianu s (Bureau 1865: 374) Bureau (1868: 274)].
Three new species were included in Tynanthus within the next years, T. igneus (Vellozo 1825: 244; 1827: tab. 15)
Rodrigues (1891: 50), T. guatemalensis Smith (1893: 6) and T. micranthus Mello ex Schumann (1894: 221), until
the genus was treated in the monograph for the Flora Brasiliensis (Bureau & Schumann 1896). In this treatment,
two new species were described, T. lindmani i Schumann (in Bureau & Schumann 1896: 409) and T. myr iant hus
Bureau & Schumman (1896: 197), totalizing eight species of Tynanthus recognized for Brazil (Bureau &
Schumann 1896). Since then, the genus has never been treated comprehensively. Instead, three new combinations
were made by various authors, i.e., T. caryophylle us (Bello 1881: 293) Alain (1965: 352), T. pa nure nsis (Bureau
1865: 373) Sandwith (1953: 465), and T. polyanthus (Bureau 1865: 378) Sandwith (1953: 465), while three others
were newly described, i.e., T. hyacinthinus Standley (1935: 87), T. macranthus Williams (1967: 250), and T.
weberbaueri Sprague (1908: 176). Furthermore, Alwyn Gentry described four new species [i.e., T. croat ianus
Gentry (1971: 93), T. pubescens Gentry (1978: 275), T. sastrei Gentry (1980: 214) and T. vil losus Gentry (1976:
60)], and transferred T. schuma nni anu s (Kuntze 1898: 243) Gentry (1974: 874) into the genus. More recently, a
synopsis of tribe Bignonieae recognized 15 species in Tynanth us (Lohmann & Taylor 2014).
Tyn anthus is clearly circumscribed and characterized by three morphological synapomorphies: a smell of
cloves on the vegetative organs, small flowers with bilabiate corollas, and fruits with raised margins (Lohmann
2006, Lohmann & Taylor 2014). The strong smell of cloves on the vegetative organs has led to the popular name
“cipó-cravo” (Brazil) or “clavo huasca” (e.g., Colombia, Peru and Ecuador). Other morphological features that are
not exclusive to Tynanthus, also being found in other genera are also useful features to recognize the genus, namely
the inflorescences in thyrses or compound thyrses, the densely pubescent corolla, thecae curved forward, densely
pubescent or velutinous ovary, and a poorly developed nectar disk (Lohmann & Taylor 2014). In addition, the
presence of trifid tendrils and lack of interpetiolar gland fields are also useful to identify species, although these
traits are variable in some species of the genus.
Although Tynanthus is strongly supported as monophyletic (Lohmann 2006, Medeiros & Lohmann 2015) and
well characterized morphologically, the delimitation of species and patterns of morphological variation within the
genus have remained unclear (Lohmann & Taylor 2014). This is in part because several of its species are only
known from few herbarium collections and also because a detailed account for the species of Tynanthus has not
been conducted since the monograph of Bureau and Schumann for the Flora Brasiliensis (1896). Extensive
herbarium studies in association with fieldwork and new molecular phylogenetic data (Medeiros & Lohmann
2015) provide new morphological, ecological, evolutionary and biogeographical information about Tynanthus, and
an excellent foundation for a new comprehensive taxonomic treatment of all species in the genus.
Phytotaxa 216 (1) © 2015 Magnolia Press
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TAXONOMIC REVISION OF TYNANTHUS (BIGNONIEAE)
Material and methods
The taxonomic treatment of Tynanthus is based on new morphological observations, and novel molecular
phylogenetic data (Medeiros & Lohmann 2015). Protologues and type collections of all species names, including
the recognized taxa and respective synonyms were examined. All accepted names are listed alphabetically, with
nomenclatural discussions and citations following McNeill et al. (2012).
Morphological characters, phenological and distributional data were collected from collections obtained
during fieldwork and specimens deposited in the following herbaria: A, BHCB, BM, BR, C, CAY, CVRD, E, ESA,
F, FUEL, G (incl. G-DC), GH, HAL, HRCB, IAC, IAN, INPA, K, LE, M, MBM, MG, MO, MPU, NY, OKL, P, R,
RB, S, SCZ, SP, SPF, SPSF, STRI, TCD, UEC, UFACPZ, UPCB, UPS, US, VEN, VIC, W (acronyms follow
Thiers, continuously updated). Morphological descriptions follow the terminology of Radford (1986), as well as
Weberling (1989) for inflorescence morphology, the Leaf Architecture Working Group (1999) for leaf venation,
Gomes-Silva (2009) for leaflet mite-domatia, Nogueira et al. (2013) for trichomes, and Lohmann & Taylor (2014)
for prophyll morphology.
A dataset with ca. 650 geo-referenced localities was compiled for all species in Tynanthus. These data were
then inserted into ArcGIS 10.2.1 (ESRI) for the preparation of distribution maps for all species recognized.
Examined specimens are ordered alphabetically, and by date within localities; question marks indicate dubious or
ambiguous information.
Results
Taxonomic treatment
Tynanthus Miers (1863: 193). Ty nna nthu s , orth. var. Lectotype (designated by Sandwith 1962a: 454): Tynanthus fasciculatus
(Vellozo) Miers.
Schizopsis Bureau (1864: 44). Type: Schizopsis labiata (Chamisso 1832: 701) Bureau (1865: 373) [= Tynanthus labiatus
(Chamisso 1832: 701) Miers (1863: 193)].
Lianas. Branchlets with four phloem wedges in cross section, strong clove odour, conspicuously tetragonal to
terete (in general, somewhat flattened when young), with or without ritidome, finely striated or not, few to densely
lenticeled (sometimes without lenticels), villous, tomentose, pubescent, puberulent or glabrescent, with simple,
peltate or patelliform trichomes; interpetiolar ridge absent or present; interpetiolar patelliform glands absent or
present; prophylls of the axillary buds minute, foliaceous or bromeliad-like, triangular, elliptic, ovate or obovate,
villous, tomentose, pubescent, puberulent or glabrescent throughout, with simple, peltate or patelliform trichomes.
Leaves 2–3 foliolated; terminal leaflets often modified into simple or trifid tendrils (rarely bifid, when very young),
with or without adhesive-disks on tips; petioles and petiolules with a more or less conspicuous canalicule on the
upper side, villous, tomentose, pubescent, puberulent or glabrescent throughout the surface or only at the upper
canalicule, with simple, peltate or patelliform trichomes; lateral petiolules with equal lengths and the terminal one
longer, when present; leaflets membranous to coriaceous, discolor or concolor, elliptic, ovate or obovate; apex
acuminate, caudate, mucronate or obtuse; base cuneate, obtuse, truncate or subcordate, symmetrical or
asymmetrical; margin entire (rarely dentate); the abaxial surface villous, tomentose, pubescent, puberulent or
glabrescent throughout or only on and near the veins, with simple, peltate or patelliform trichomes; the adaxial
surface villous, tomentose, pubescent, puberulent, or glabrescent throughout or only on and near the veins, with
simple, peltate or patelliform trichomes; glandular trichomes evenly distributed throughout both surfaces or
especially on one surface; first venation pinnate; second venation weak brochidodromous or brochidodromous;
third venation alternate percurrent (sometimes random reticulate); pocket domatia with or without trichomes.
Inflorescence axilar or terminal, a thyrse or a compound thyrse, lax, with conical aspect, or dense, with corymbose,
subcorymbose or conical aspect; axis villous, tomentose or pubescent, with simple, peltate or patelliform
trichomes; bracts of the inflorescence caducous or persistent, triangular to linear triangular, villous to pubescent or
glabrescent throughout or only at margins; floral bracts triangular. Calyx green to yellowish, grayish or reddish,
campanulate, membranous to chartaceous, with transversal or oblique aperture, truncate, denticulate or laciniate,
MEDEIROS & LOHMANN
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Phytotaxa 216 (1) © 2015 Magnolia Press
tomentose, pubescent, puberulent or glabrescent outside, with simple and peltate trichomes, with or without
patelliform glands, glabrous inside. Corolla white, cream or pale yellow (sometimes pale lilac, pale green, pale
pink, pale red or pale blue), bilabiate, with two (almost totally fused) upper lobes and three lower lobes, densely
pubescent outside, with simple and peltate trichomes, internally glabrous at the top of the tube, tomentose to
pubescent or glabrescent at the base, with simple and/or long and short stipitate trichomes; nectar guides present or
absent, with long and short stipitate trichomes; lobes entire, densely pubescent to pubescent throughout or only at
margins, with simple and peltate trichomes, acute, obtuse or rounded. Androecium with fertile stamens inserted at
the same position; filaments with long and short stipitate trichomes at the base; anthers thecae cream, obovate to
elliptic, divergent and reflexed forward, glabrous, inserted or subexserted; staminode glabrous, glabrescent or with
long and short stipitate trichomes. Gynoecium with ovary conical to oblong, densely pubescent or velutinous, with
simple trichomes, with a ring of longer trichomes at the base, with 2 or 4 series of ovules per locule; nectar disc not
evident; style tomentose to pubescent at the base, with simple trichomes; stigma with lamellae lanceolate, glabrous.
Fruit a linear flattened to subtetragonal capsule, with extremities acuminate, acute or obtuse, coriaceous to woody,
smooth or granular throughout, without lenticels to densely lenticeled, villous, tomentose, pubescent, puberulent or
glabrescent, with simple, peltate or patelliform trichomes; central ridge double or single, prominent or not; margins
slightly or prominently raised. Seeds thin, bialatae, more or less oblong, finely striated; body brown; wings hyaline-
membranaceous, sharply demarcated from the body.
Key to species of Tynant hus
1 Lax thyrses, with conical aspect............................................................................................................................................. 2
- Dense thyrses, with corymbose or subcorymbose aspect (or conical in T. d en siflor us ).......................................................11
2 Tendrils simple ....................................................................................................................................................................... 3
- Tendrils trifid (rarely bifid, in young individuals of T. co gna tu s).......................................................................................... 4
3 Leaflets with apex caudate-mucronate; calyx with patelliform glands; fruits winged, with margins prominently raised ......
....................................................................................................................................................................... T. gu at emal en sis
- Leaflets with apex acuminate-mucronate; calyx without patelliform glands; fruits unwinged, with margins slightly raised
............................................................................................................................................................................ T. pol ya nthus
4 Prophylls foliaceous............................................................................................................................................................... 5
- Prophylls minute, triangular to shallowly triangular, or bromeliad-like................................................................................ 6
5 Leaflets discolor; corolla 1.2–1.7 cm long......................................................................................................... T. p anur ensi s
- Leaflets concolor; corolla 0.6–0.8 cm long............................................................................................................... T. s ast rei
6 Prophylls bromeliad-like........................................................................................................................................................ 7
- Prophylls minute, triangular to shallowly triangular.............................................................................................................. 8
7 Leaflet domatia pubescent; inflorescence axis without patelliform trichomes; Atlantic forest, Brazil (SE)...........................
....................................................................................................................................................................T. espiritosantensis
- Leaflet domatia glabrous; inflorescence axis with patelliform trichomes; Amazon forest, Bolivia, Brazil (N) and Peru......
..................................................................................................................................................................... T. sc hu mann ia nus
8 Interpetiollar patelliform glands present; fruits with double central ridge ......................................................... T. pub es cens
- Interpetiolar patelliform glands absent; fruits with single central ridge ................................................................................ 9
9 Branchlets tomentose to pubescent throughout; calyx without patelliform glands; fruits unwinged, with margins slightly
raised ..................................................................................................................................................................... T. cog na tus
- Branchlets glabrescent (if pubescent, only at nodes); calyx with patelliform glands; fruits winged, with margins promi-
nently raised ......................................................................................................................................................................... 10
10 Petioles and petiolules with patelliform trichomes; corolla 1–1.4 cm long, with nectar guides............................. T. labi at us
- Petioles and petiolules without patelliform trichomes; corolla 0.5–0.9 cm long, without nectar guides .......... T. m ic rant hu s
11 Interpe tiolar patelliform glands present ............ ................... .............. ....................................... ........................................... 12
- Interpetiolar patelliform glands absent................................................................................................................................. 13
12 Tendrils trifid; corolla 0.8–1.5 cm long.............................................................................................................. T. d en sifl or us
- Tendrils simple; corolla 2–3.8 cm long............................................................................................................. T. ma cr anth us
13 Young branchlets pubescent; calyx minutely denticulate (sometimes truncate); fruits unwinged, with margins slightly
raised ................................................................................................................................................................... T. cro atian us
- Young branchlets tomentose; calyx laciniate; fruits winged, with margins prominently raised...................... T. fascic ul atus
Phytotaxa 216 (1) © 2015 Magnolia Press
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TAXONOMIC REVISION OF TYNANTHUS (BIGNONIEAE)
1. Tynanthus cognatus (Cham.) Miers (1863: 193). Bignonia cognata Chamisso (1832: 703). Lectotype (designated
here):—BRAZIL. "Brasil aequinoct.", s.d., F. Sellow s.n. (US! (barcode 125821)).
Tyn an thus e lega ns Miers (1863: 193). Bignonia elegans Chamisso (1832: 702), nom.illeg., non Bignonia elegans Vel l oz o 1 82 5 ;
1827. Lectotype (designated here):—BRAZIL. Sin loc., s. d., F. S e l l o w s . n . (US! (barcode 125825)). syn. nov.
Schizopsis chimonantha Bureau (1865: 375). Lectotype (designated here):BRAZIL. Bahia:Prope Ilheos, 1838, B.
Luschnath s.n. (BR!; isolectotype BR!). Syntype: BRAZIL. Rio de Janeiro: “Bords de la rivière d’Hytu, près la fazenda de
Bemfica”, 1816–1821, A. St.-Hilaire Catal. D, Nº 25 (P! (barcodes 3606733, 3606734), K!).
Schizopsis regnelliana Bureau (1865: 376). Lectotype (designated here):—BRAZIL. Minas Gerais: “Caldas, 28 December
1859, A. F. Regnel III-52 (S (14-19845) photo!; isolectotype K!). Sy ntypes:—BRAZIL. São Paulo: “Prés de la
Paranapitanya”, 1816–1821, A. St.-Hilaire Catal. C2, Nº 1342 (P! (barcodes 3606735, 3606736)). Minas Gerais: Sin. loc.,
1845, Widg ren 7 43 (BR!).
Fig. 1: A–E
Lianas. Branchlets tetragonal to terete, with or without ritidome, finely striated, lenticeled to densely lenticeled,
tomentose to pubescent, with simple and peltate trichomes; interpetiolar ridge present (sometimes absent);
interpetiolar patelliform glands absent; prophylls of the axillary buds 0.7–1.7 mm long, 0.6–1.5 mm wide, minute,
shallowly triangular to triangular, tomentose to pubescent or glabrescent throughout, with simple and peltate
trichomes. Leaves 2–3 foliolated; terminal leaflets often modified into trifid tendrils (rarely bifid, when very
young), sometimes with adhesive-disks on tips; petioles and petiolules tomentose to puberulent throughout the
surface, with simple and peltate trichomes; petioles (0.2–)0.5–3.6 cm long; petiolules (0.3–)0.5–2.3 cm long;
leaflets (1.6–)4–10.6 cm long, (0.6–)1.8–6.2 cm wide, chartaceous to coriaceous, discolor, obovate to elliptic; apex
acuminate or obtuse, mucronate; base cuneate or obtuse, symmetrical; margin entire; the abaxial surface tomentose
to pubescent throughout (sometimes only on and near the veins), with simple, peltate and patelliform trichomes;
the adaxial surface pubescent to glabrescent throughout or only on and near the veins, with simple, peltate and
patelliform trichomes; glandular trichomes evenly distributed throughout both surfaces; second venation weak
brochidodromous or brochidodromous; pocket domatia with (sometimes without) trichomes. Inflorescence axilar
or terminal, a thyrse or a compound thyrse, lax, with conical aspect, first order (2.5–)3–13 cm long, second order
2.5–5.5 cm long; axis tomentose to densely puberulent, with simple and peltate trichomes; bracts of the
inflorescence predominantly caducous, tomentose to pubescent throughout, 0.5–1.9 mm long; floral bracts 0.3–0.7
mm long; floral pedicels 1–7.5 mm long. Calyx green, 1.3–2.7 mm long, 1.3–2.3 mm wide, with transversal
(sometimes oblique) aperture, truncate or minutely 5-denticulate, tomentose to puberulent throughout outside,
without patelliform glands; lobes 0.1–0.4 mm long. Corolla white, cream or pale yellow (sometimes pale lilac),
0.5–1.1 cm long, 2.3–3.7 mm wide at the tube opening; tube 3–4.9 mm long, internally pubescent to glabrescent at
the base, with simple and long and short stipitate trichomes; nectar guides absent, but with a path of long and short
stipitate trichomes; lobes densely pubescent to pubescent throughout lower ones and at the margin of upper ones;
upper ones 0.3–1.7 mm long, 0.6–1.9 mm wide, acute to obtuse; lower ones 1.6–3.6 mm long, 1.2–3.1 mm wide,
obtuse to rounded. Androecium with fertile stamens inserted 1–1.5 mm from the base of the corolla; shorter ones
2.5–3.5 mm long; longer ones 3.5–4.5 mm long; anthers thecae 0.7–0.8 mm long, obovate to elliptic, subexserted;
connective extending 0.1–0.2 mm beyond anther attachment; staminode 1.3–1.7 mm long, glabrous. Gynoecium
4.5–8.9 mm long; ovary 1–1.3 mm long, 0.7–0.9 mm wide, conical, velutinous; style 3.5–7.5 mm long, tomentose
at the base. Fruit a linear flattened capsule, (6.5–)10–25 cm long, 0.5–1.1 cm wide, coriaceous, smooth to granular
near the midvein and granular near the margins, without lenticels to densely lenticeled, tomentose to pubescent,
with simple and peltate trichomes; central ridge single, slightly or not prominent; margins slightly raised
(unwinged), 0.1–0.2 cm wide. Seeds body (0.7–)0.9–1.7 cm long, (0.3–)0.5–0.7 cm wide; wings (0.3–)0.5–1 cm
long.
Phenology:—Flowers from October to March and produces fruits from February to September.
Distribution and habitat:—Occurs in moist broadleaf forests of Brazil (Bahia, Espírito Santo, Minas Gerais,
Paraná, Rio de Janeiro, Rio Grande do Sul, Santa Catarina and São Paulo) (Fig. 2).
Additional specimens examined:—BRAZIL. Sin loc., s.d., A. Glaziou s.n. (F 539388). Sin loc., s.d., J.C.
Mello 8yt (P barcode 3606711). Sin loc., s.d., A.F. Regnell s.n. (K). Sin loc., s.d., L. Riedel s.n. (NY). Sin loc.,
s.d., F. Se l l o w 1 0 74 (BM). Sin loc., 1814–1817, J. Bowie & A. Cunningham s.n. (BM). Sin loc., 1823, L. Riedel?
s.n. (LE). Bahia: Rio Gongogi Basin, 100–500 m, 1 October–30 November 1915, H.M. Curran 213 (US);
Rodovia Itabuna–Camacan, 12 km L de Itabuna, 8 April 1965, R.P. Belém & M. Magalhães 719 (IAN, K, NY).
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FIGURE 1. A–E. Tynanthus cognatus: A. Flowering branch; B. Open corolla showing the androecium; C. Gynoecium (D.A. Folli
1795, SPF); D. Fruit; E. Seed (J.S. Carneiro 161, FUEL). F–N. T. c roa ti an us: F. Flowering branch; G. Leaflet; H. Flower; I–J. Calyx
with patelliform glands; K–M. Stamen with stipitate trichomes at the base of filaments and curved thecae (S. Knapp 1053, NY); N.
Fruit (A.H. Gentry 6696, NY). Illustration by Klei Sousa.
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TAXONOMIC REVISION OF TYNANTHUS (BIGNONIEAE)
Rodovia Banco Central a Gongogi, 17 March 1971, Raimundo S.P. 1163 (RB). Ilhéus, Área do CEPEC (Centro de
Pesquisas do Cacau), km 22 da Rodovia Ilhéus–Itabuna (BR-415)?, 50 m, 17 February 1982, J.L. Hage 1641 (K,
MO); CEPLAC, Quadra “D”, Matinha das Preguiças, 1 August 2007, R.O. Perdiz et al. 84 (SPF). Jussari, RPPN
Serra Teimoso, entrada 7.5 km da Rodovia Jussari–Palmira, Fazenda Teimoso, 1.7 km da entrada, 26 January 2006,
J.L. Paixão et al. 684 (SPF). Maraú, 13 January 1967, R.P. Belém & R.S. Pinheiro 3129 (K, MO, NY). Una,
Rodovia São José–Una, ca. 9 km da rodovia BR-101, 18 March 1999, J.G. Jardim et al. 2063 (SPF). Espírito
Santo: Conceição do Castelo, Alto Bananal, 6 November 1986, G. Hatschbach & J.M. Silva 50691 (MBM, MO).
Itapemerim, Fazenda do Ouvidor, Usina Paineiras, acesso pela Rodovia ES-490, entrada à esquerda, 2.5 km após o
trevo da SAFRA, em direção a Marataízes, 29 December 2007, A.M. Assis et al. 1292 (SPF). Linhares, Reserva
Natural da Companhia Vale do Rio Doce, Estrada 243, 31 January 1985, A. Peixoto et al. 3037 (MO); Ibid., 2
February 1985, A. Peixoto et al. 3404 (MO, NY); Ibid., 2 February 1985, A. Peixoto et al. 3406 (F photo, K, MO,
NY); Ibid., 20 m, 14 September 1987, A.H. Gentry et al. 59208 (MO); Ibid., Aceiro com a LASA, próximo à
estrada Gonçalo Alves, 18 January 1993, D.A. Folli 1795 (CVRD, SPF, US); Ibid., Próximo ao aceiro com a Lasa,
14 June 1993, D.A. Folli 1898 (CVRD, SPF); Ibid., 150 m, 15 July 2001, L.G. Lohmann & B. Whitney 663 (CVRD,
MO); Ibid., at the end of the road “Peroba Amarela”, 150 m, 17 July 2001, L.G. Lohmann et al. 680 (CVRD, MO);
Ibid., Aceiro com Catelã Jueirana, Estrada Orelha de Macaco, km 2.4, 17 December 2001, D.A. Folli 4148 (CVRD,
SPF); Ibid., Estrada Municipal do Canto Grande, próximo à entrada da cabana Martinelli, 32 m, 2 May 2008, A.R.
Zuntini et al. 241 (CVRD, MO, RB); Ibid., Estrada Municipal Canto Grande, 36 m, 6 January 2009, D.A. Folli
6274 (CVRD); Ibid., Estrada Flamengo, após o cruzamento com a Gonçalo Alves, 40 m, 28 January 2014, M.C.
Medeiros & R.B. Louzada 42 (CVRD, SPF); próximo à Reserva Natural da Companhia Vale do Rio Doce,
Projeto1, córrego da Jacutinga, APP da Aracruz, 37 m, 2 May 2008, A.R. Zuntini et al. 240 (MO, RB). Pinheiros,
Reserva Biológica Córrego do Veado, 9 May 2008, D.A. Folli 6021 (CVRD). Santa Leopoldina, Distrito de
Mangaraí, Cachoeira do Retiro, 2 October 2005, M.O.S. Crepaldi 54 (RB). Santa Teresa, Vale do Canaã, 800–850
m, 1 February 1969, D. Sucre & P.I.S. Braga 4576 (MO, RB); Várzea Alegre, Cachoeira do Magdalon, 26 October
2000, V. Demuner et al. 1510 (SPF). Sooretama, Estrada Municipal Canto Grande, próximo à Reserva Natural da
Companhia Vale do Rio Doce (“Reserva de Linhares”), 34 m, 13 December 2007, A.R. Zuntini et al. 147 (CVRD,
SPF). Minas Gerais: Sin loc., 1845, Widgren s.n. (BR, P barcode 3606716). Sin loc., Campos da Mantiqueira,
perto do vale do Rio Verde, January 1885, J. Saldanha s.n. (R 128076). Caldas, 1868, S. Henschen s.n. (US
201385). Coronel Pacheco, 3 March 1943, E.P. Heringer 1176 (RB); Água Limpa, 10 December 1946, E.P.
Heringer 2519 (SP). Faria Lemos, Fazenda Santa Rita, 600 m, 6 February 2000, L.S. Leoni & A.E. Silva 4366
(MO). Monte Belo, Fazenda Lagoa, 800 m, 6 September 1987, A.H. Gentry et al. 59095 (MO, UEC); Ibid., 6
September 1987, A.H. Gentry et al. 59101 (MO, UEC); Ibid., 6 September 1987, M.C.W. Vieira 1235 (RB). Poços
de Caldas, Fazenda Chiqueirão, 3 December 1981, H.F. Leitão Filho et al. 1583 (BHCB, FUEL, UEC). Tombos,
Fazenda da Cachoeira, 12 July 1935, M. Barreto 1563 (MO, R). Paraná: Antonina, Sapitanduva, 18 January 1974,
G. H a t s c h b a c h 3 3 6 7 1 (M, MO, SPF, UEC, US). Apucarana, Parque da Raposa, 22 February 2005, J.S. Carneiro et
al. 297 (FUEL). Arapongas, Fazenda do Bule, 22 June 1999, D.A. Estevan et al. 89 (FUEL, VIC). Cerro Azul,
Cabeceira do Ribeirão do Tigre, 8 December 1983, G. Hatschbach 48829 (MO, US). Fênix, Parque Estadual Vila
Rica do Espírito Santo, 11 December 1998, M. Borgo & S.M. Silva 349 (NY, UPCB). Ibiporã, Chácara Alcides
Pelisson, Água Tucano, 28 May 1989, A. Pelisson & F.M.E. Longhi s.n. (FUEL 7382); Fazenda Doralice, 1
December 1995, M.C. Dias et al. 21 (FUEL). Jaguariaíva, 740 m, 13 January 1915, P. D u s é n 1 6 3 4 1 (F, K, NY).
Jundiaí do Sul, Fazenda Monte Verde, 16 December 1999, J. Carneiro 848 (SPF); Ibid., Mata do Cruzeiro, 14
January 2000, G. Hatschbach et al. 69944 (SPF); Mata do Cruzeiro, 3 March 2003, J. Carneiro 1393 (MBM).
Londrina, Parque Municipal Arthur Thomas, 19 December 1984, M.I.O.J. Neves et al. s.n. (FUEL 523); Ibid., 13
March 1986, A.C. Amorin s.n. (MO 3386506); Ibid., Trilha da Capivara, 520 m, 13 October 2011, M.C. Medeiros
& E.F. Rossetto 33 (SPF); Floresta dos Irmãos Godoy, 21 August 1985, L.A.C. Rodas et al. 8 (FUEL); Ibid., 28
November 1985, F.C. Silva et al. 950 (FUEL); Ibid., 20 March 1986, L.N. Pizzaia et al. 45 (FUEL, MO); Ibid., 14
January 1989, L.H.S. Silva & F.C. Silva 170 (FUEL, K, MBM, UPCB); Ibid., 9 July 1997, R. Irío & L. Lima 194
(UFACPZ); Ibid., 16 July 1997, V. F. K in u p p et a l . 1 6 30 (FUEL); Mata do IAPAR, 30 June 1988, L.A. Volpato s.n.
(FUEL 6375, HRCB 34681); Paiquerê, 6 February 1997, V.F. Kinupp et al. 230 (FUEL); Fazenda Figueira-
Paiquerê, Fragmento 16, 19 February 2003, M.C. Lovato et al. 388 (FUEL, R); Ibid., Fragmento 16, 19 February
2003, M.C. Lovato et al. 400 (ESA, FUEL, MBM); Ibid., 19 February 2003, D.A. Estevan s.n. (HRCB 44200,
44201); Ibid., 2004, J.S. Carneiro et al. 164 (FUEL, HRCB); Ibid., Fragmento 2, 2 April 2004, J.S. Carneiro et al.
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162 (FUEL); Ibid., Fragmento 2, 13 April 2004, J.S. Carneiro 163 (FUEL, R); Ibid., Fragmento 19, 15 June 2004,
J.S. Carneiro et al. 161 (FUEL); Ibid., Fragmento 2, 22 June 2004, J.S. Carneiro et al. 173 (FUEL); Ibid.,
Fragmento 1, 1 July 2004, J.S. Carneiro et al. 167 (FUEL). Ortigueira, Futuro eixo da Barragem, margem esquerda
do Rio Tibagi, 30 October 2008, M. Kaehler 338 (MBM, UPCB). Ribeirão do Pinhal, Fazenda São Pedro, 11
February 2001, J. Carneiro 1068 (SPF). Rolândia, December 1936, G. T e s s m a n n 6 0 0 1 (MBM, SP). São Jerônimo
da Serra, 22 December 1999, C. Medri et al. 305? (FUEL 29356). São Pedro do Ivaí, 23 January 1991, F. B a r r o s
2112 (SP); Fazenda Santa Bárbara?, 18 December 2003, O.S. Ribas et al. 5668 (G, K, MBM, MO, RB, SPF).
Telêmaco Borba, Rio Tibagi, próximo à ponte, 12 December 1996, V.F. Kinupp et al. 75 (FUEL, HRCB, SPF);
Margem da estrada de acesso ao eixo da Barragem, 680 m, 22 September 2008, M. Kaehler 276 (MBM). Tereza
Cristina (“Theresina”), 27 November 1911, P. D u sé n 1 11 7 2 (BM, K, US). Ventania, Fazenda Santo Expedito, 4
May 2004, D.A. Estevan et al. 498 (FUEL). Rio de Janeiro: Serra Tingua, 1780, H.W. Schott 5971 (F). Sin loc.,
1830, L. Riedel s.n. (BM, G, K, P barcode 3606732). Sin loc., 18311833, Gaudichaud 559? (P barcode 3606674).
Sin loc., 1832, L. Riedel 231 (LE). Fazenda do Sobral, 8 September 1881, A. Glaziou et al. 12973 (LE, P). Pr.
Sumidouro, na Rodovia BR-3, entre Itaipava e Pedro do Rio, 600 m, 10 December 1956, G.F.J. Pabst 10319 (RB).
E of Rio Bonito, between Niteroi and Silva Jardim, 100 m, 19 January 1985, A.H. Gentry & E. Zardini 49709
(MO); Ibid., A.H. Gentry & E. Zardini 49727 (MO). Cabo Frio, Parque Ecológico Municipal do Mico-Leão-
Dourado, 13 June 2003, G. S. Z . R e z e n de e t a l . 1 6 2 (RB); Casemiro de Abreu, Monte São João, 3 February 1970,
S.P.S. s.n. (RB barcode 58285). Guapimirim, Parque Nacional da Serra dos Órgãos, trilha do Poço da Preguiça, 410
m, 6 January 2011, M.C. Medeiros et al. 28 (SPF). Nova Iguaçu (“Iguassú”), s.d., Sin col. 8061 (P barcode
3606667). Petrópolis, Mandiocca, 1821–1824, L. Riedel s.n. (MO 4618844, NY barcode 483757, NY barcode
1032824); February–March 1823, L. Riedel s.n. (LE). Rio de Janeiro, Floresta da Tijuca, Gávea, Caminho do
Macaco, 1865?, A. Glaziou 2638 (BR, C photo in F, K, P); 8 March 1871, A. Glaziou 4719 (P, US); Ibid.,
Corcovado, 22 May 1870, A. Glaziou 4124 (P); 22 May 1870, A. Glaziou 4683 (P, US); Ibid., Grande cascade, 24
June 1870, A. Glaziou 4692 (P); Ibid., 21 January 1871, A. Glaziou 4709 (BM, K, NY, P, US); Ibid., Estrada de
Sumare, km 5, 300 m, 19 January 1975, W. B en so n 45 (MO); Prope hort. bot., 29 November 1888, W. Schwacke
s.n. (K, R 23784); Mundo Novo, Botafogo, May 1921, J.G. Kuhlmann s.n. (R 23803; RB barcode 58297); Gávea,
Mesa do Imperador, 10 March 1950, J.G. Kuhlmann s.n. (RB barcode 58293; RB barcode 58341; NY barcode
483758); Jardim Botânico do Rio de Janeiro, orla do parque, 13 September 1991, A.F. Vaz et al. 949 (RB). Santa
Maria Madalena, Mata do Laureano Vicente, 7 March 1934, S. Lima & Brade 13194 (R, RB). Teresópolis, Próximo
à Fazenda Boa Vista, 12 January 1943, H.P. Velloso s.n. (MO 2286403). Rio Grande do Sul: Vila Manresa p.
Porto Alegre, 25 July 1949, B. Rambo 42705 (MO). Caxias do Sul, Santa Lucia do Piaí, 780 m, 27 January 1999, A.
Kegler 158 (M, MBM, US). Vale do Sol, Linha XV de Novembro, 23 January 1993, J.A. Jarenkow & D.B.
Falkenberg 2281 (MBM). Santa Catarina: Sin loc., June 1868, F. M ü l l e r 1 6 6 (K). Blumenau, Bom Retiro, Mata
da Companhia Hering, 350 m, 17 September 1959, P. R . R e i t z & R . M . K l ei n 9 1 0 2 (K, US). Florianópolis, Morro
Costa da Lagoa, 300 m, 15 February 1967, R.M. Klein 7238 (K, R); Morro da Cutia, Tapera, Ribeirão, 150 m, 20
January 1970, R.M. Klein & Bresolin 8542 (K, R). Itajaí, s.d., F. M ü l l e r 2 9 8 (R). Jacinto Machado, Sanga da Areia,
200 m, 13 July 1959, P. R. R e i tz & R . M. K l e in 8 9 36 (K, US); Ibid., 250 m, 27 January 1960, P.R. Reitz & R.M. Klein
9424 (BR, F photo, G, K, M, NY, UPCB, US). Lauro Müller, Novo Horizonte, 450 m, 15 January 1959, P. R . R e i t z
& R.M. Klein 8245 (BR, F, G, K, M, NY, SP, US). Luis Alves, Braço Joaquim, 250 m, 13 January 1955, R.M. Klein
1054 (K, NY, US). Palhoça, Morro do Cambirela, 300 m, 18 January 1972, R.M. Klein & Bresolin 10008 (K). Rio
do Sul, 350 m, 31 December 1958, P. R . R e i t z 6 14 7 (BR, K, NY, US); Serra do Matador, 550 m, 12 March 1959,
P. R . R e i t z & R . M . K l e i n 8 5 3 7 (K, US). São Bento do Sul, Braço esquerdo, 417 m, 21 November 2009, T.J. Cadorin
et al. 799 (SPF). São Paulo: Sin loc., 27 November 1871, J.C. Mello 51 (US). Sin loc., 25 December 1873, H.
Mosén 1488 (P). Barretos, Margem do Rio Pardo, November 1917, A. Sampaio s.n. (R 23545, RB barcode 66737).
Campinas, Barão Geraldo, Santa Genebra Forest Reserve, Transect 3, 550 m, 27 August 1987, A.H. Gentry & A.
Silva 58727 (MO, UEC); Ibid., Transect 4, 550 m, 27 August 1987, A.H. Gentry 58752 (MO, UEC). Ipeúna,
Ribeirão Passa-Cinco, 26 January 1984, A. Furlan 175 (HRCB). Marília, Estação Experimental, 14 January 1993,
G. D ur i g a n 30 6 91 (UEC). Rio Claro, Fazenda São José, 15 November 2000, R. Ubulutsch & M.A. Assis 101
(HRCB); Ibid., trilha que atravessa o fragmento, 20 December 2000, R. Udulutsch & M.A. Assis 134 (HRCB). São
Paulo, Bosque da Avenida, 17 December 1933, J.G. Kuhlmann s.n. (RB barcode 58318); Jardim Botânico, 15
January 1938, O. Handro s.n. (SP 43041); Ibid., Trilha Fontes do Ipiranga, 24 October 2006, B.L.P. Villagra 225
(SP); Cidade Jardim, 4 January 1944, W. Hoehne 1184 (MO, SPF); Serra da Cantareira, Picada Dom Bento, Pedra
Grande, 1000 m, 6 January 1953, F. M a r k g ra f s . n. (SPSF 4061).
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TAXONOMIC REVISION OF TYNANTHUS (BIGNONIEAE)
Taxonomic notes:—The morphological similarity between Tynanthus cognatus and T. elegans has long been
noted. Indeed, Chamisso (1832) commented on their overall similarity when those taxa were originally described
as Bignonia cognata and B. elegans, respectively. On the other hand, Chamisso (1832) also noted differences in the
type and density of the indumentum encountered in the vegetative and reproductive organs of these taxa, which he
considered sufficient to maintain the taxa as separate. However, a detailed analysis of the indumentum of these
species along the entire range of distribution of these taxa showed that indumentum density varies geographically.
More specifically, the collections from the northern portion of the range (states of Bahia and Espírito Santo) were
shown to have glabrescent branches and leaves, while the specimens from the southern portion (states of Paraná
and Rio Grande do Sul) are generally densely pubescent to pubescent. Therefore, indumentum variation is not a
diagnostic trait and not sufficient to keep those taxa as separate. Furthermore, molecular phylogenetic data
(Medeiros & Lohmann 2015) indicated that T. elegans is nested within T. cognatus, corroborating the
synonimization of T. elegans in T. cognatus.
FIGURE 2. Distribution of Tyn ant hu s co gna tu s.
Tynanthus cognatus can be easily recognized by its obovate to elliptic leaflets. Tynanthus pubescens is the
only other species of Tynant hus to present this feature; however, these species can be separated by the winged fruits
of T. pubescens (versus unwinged in T. c ogn atus) and Amazonian distribution (versus Atlantic Forest in T.
cognatus). Tynanthus cognatus shares lax inflorescences and unwinged fruits with its sister species T. polyanthus
(Medeiros & Lohmann 2015). Nevertheless, T. cognatus is easily distinguished from T. polyanthus by the minute
prophylls (versus foliaceous prophylls in T. pol yanthus) and the inconspicuously tetragonal young branchlets
(versus conspicuously tetragonal in T. polyanthus).
Nomenclatural notes:—Chamisso (1832) described T. cognatus and T. elegans based on Sellow’s collections.
Chamisso’s types are deposited at LE (Stafleu & Cowan 1976: 482), but we were not able to locate those materials
during two visits to the LE herbarium. The types of T. cognatus and T. elegans deposited at B were destroyed
during the World War II (Hiepko 1987) and are no longer available. Despite that, we were able to access photos of
the B types from the F website. The photographed collection of T. cognatus was labelled as Sellow 166 while the
photographed collection of T. e legans was labelled Sellow 5596. Unfortunately, we were unable to locate any
duplicates of the Sellow 166 and 5596 collections in any of the herbaria visited. Instead, we were able to locate a
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Sellow unnumbered collection at K that matches the identifications of T. cognatus (K barcode 449541). In addition,
we were also able to locate two Sellow unnumbered collections at US that had been correctly identified as
Bignonia cognata (US barcode 125821) and B. elegans (US barcode 125825), respectively; these materials are here
selected as lectotypes. The calligraphy of the collection US barcode 125821 matches Chamisso's hand writing. The
identification of the US barcode 125825 collection was made by Schumann, who worked at B and accessed
Chamisso's types kept in that herbarium.
Two syn type s we re cited by Bureau (1 865) in the prot ologue of Schizopsis chimonantha (Luschnath s.n. and
St. Hilaire Catal. D N
o
25) and four syntypes in the protologue of S. regnelliana (Regnel III-52, St. Hilaire Catal.
C2 N
o
1342, Widgren 743, and Martius s.n.). For S. chimonantha, four sheets of the collection Luschnath s.n. were
located at BR. The duplicate that has a hand-written label with calligraphy that matches Bureau’s hand writing,
with the identification of Tynanthus cognatus is here selected as a lectotype; this material also represents the best
quality material from the four duplicates. For S. regnelliana, three duplicates of the Regnell III-52 were located in
the S herbarium (cited in the protologue), one dated from 1859 (S 14-19845), one from 1867 (S 14-19860) and one
from 1868 (S 14-19863). The only material whose date predates the publication of the protologue of S. regnelliana
(S 14-19845) is here selected as a lectotype. Additional duplicates of the Regnell III-52 collection were located in
other herbaria (i.e., K!, LE!, MO!, P!, R!, US!); however, Regnell’s collection numbering system often reflect
individual species and not collections from a single individual and thus, do not represent duplicates of the holotype.
We chose one of the Regnell’s collections as lectotype because of the good conditions of the material and because
the epithet “regnelliana” suggests that this material was described based on a Regnell collection. It is also important
to note that a Martius collection, presumably kept at M, was also cited in the protologue of S. regnelliana.
Unfortunately, we were unable to locate this material. Furthermore, this collection is also cited in the protologue of
Arrabidaea fasciculata [= T. fasciculatus (Vell.) Miers], complicating its identity. This material was thus excluded
from the list of syntypes of T. regnelliana.
2. Tynanthus croatianus Gentry (1971: 93) (as “Tynnanthus”). Type:—PANAMA. Panama: Shoreline of broad-
mouthed cove NE of Drayton House on Barro Colorado Island, 28 August 1970, T.B. Croat 11927 (holotype MO!
(2016962); isotypes F! (1697804), GH! (barcode 93264) photo, K! (barcode 449552), MO! (2039180, 2042197,
2042198), NY! (barcode 328978), SCZ! (barcode 4087) photo, STRI! (759) photo, US! (barcode 125784)).
Fig. 1: F–N
Lianas. Branchlets tetragonal to terete, without ritidome, finely striated, lenticeled to densely lenticeled, pubescent
to glabrescent, with simple and peltate trichomes; interpetiolar ridge absent (sometimes present); interpetiolar
patelliform glands absent; prophylls of the axillary buds 1.3–2.5 mm long, 1–2.5 mm wide, minute, shallowly
triangular, pubescent or puberulent to glabrescent throughout, with simple and peltate trichomes. Leaves 2–3
foliolated (more commonly 2); terminal leaflets often modified into trifid tendrils, without adhesive-disks on tip;
petioles and petiolules pubescent throughout surface or only at the upper canalicule, with simple, peltate and
patelliform trichomes; petioles (1.4–)1.8–7.5 cm long; petiolules (0.5–)1.1–3.8 cm long; leaflets (3.5–)5–11.4 cm
long, (1.4–)3–9 cm wide, membranous to chartaceous (sometimes subcoriaceous), discolor, ovate to elliptic; apex
acuminate or caudate, mucronate; base cuneate, obtuse, truncate or subcordate, symmetrical or asymmetrical;
margin entire (rarely dentate); the abaxial surface pubescent to glabrescent on and near the veins (sometimes
throughout), with simple, peltate and patelliform trichomes; the adaxial surface pubescent to glabrescent
throughout or only on and near the veins, with simple, peltate and patelliform trichomes; glandular trichomes
evenly distributed throughout both surfaces; second venation weak brochidodromous; pocket domatia without
trichomes. Inflorescence axilar or terminal, a thyrse, dense, with corymbose or subcorymbose aspect, (2.3–)3.3–6.5
cm long; axis densely pubescent to pubescent, with simple, peltate and patelliform trichomes; bracts of the
inflorescence predominantly caducous, densely pubescent to pubescent throughout, 0.5–1.5 mm long; floral bracts
0.4–0.6 mm long; floral pedicels 1–5 mm long. Calyx green, 3–4 mm long, 3–4 mm wide, with transversal
aperture, minutely 5-denticulate (sometimes truncate), densely pubescent to puberulent throughout outside, with
patelliform glands; lobes 0.1–0.3 mm long. Corolla white, 1.2–2 cm long, 4–7 mm wide at the tube opening; tube
5–9 mm long, internally tomentose at the base, with simple and long and short stipitate trichomes; nectar guides
present, yellow; lobes densely pubescent to pubescent throughout lower ones and at the margin of upper ones;
upper ones 1–3 mm long, 1–4 mm wide, acute to obtuse; lower ones 3–7 mm long, 3–5.5 mm wide, obtuse to
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TAXONOMIC REVISION OF TYNANTHUS (BIGNONIEAE)
rounded. Androecium with fertile stamens inserted ca. 2 mm from the base of the corolla; shorter ones 7–9(–10)
mm long; longer ones 8–10(–12) mm long; anthers thecae 1.4–1.9 mm long, obovate to elliptic, subexserted;
connective extending 0.2–0.3 mm beyond anther attachment; staminode 4–5 mm long, glabrous. Gynoecium
13–15(–17) mm long; ovary 1.5–2 mm long, 0.8–1.1 mm wide, conical, densely pubescent; style 11–13 mm long,
tomentose to pubescent at the base. Fruit a linear flattened capsule, 16–37 cm long, 0.7–1.2 cm wide, coriaceous to
woody, smooth to granular near the midvein and granular near the margins, without lenticels to densely lenticeled,
densely pubescent to pubescent, with simple and peltate trichomes; central ridge single, slightly or not prominent;
margins slightly raised (unwinged), 0.1–0.3 cm wide. Seeds body 1.1–1.9 cm long, 0.4–0.8 cm wide; wings 0.6–1.1
cm long.
Phenology:—Flowers from July to September and fruits from October to March.
Distribution and habitat:—Occurs in moist broadleaf forests from Colombia (Chocó), Costa Rica
(Puntarenas) and Panama (Colón, Darién and Panamá) (Fig. 3).
FIGURE 3. Distribution of Tynanthus croatianus.
Additional specimens examined:—COLOMBIA. Chocó: Trail from Unguia along Rio Tigre toward base of
Serrania del Darién, 200–300 m, 16 July 1975, A.H. Gentry & L.E. Aguirre 15209 (MO). COSTA RICA.
Puntarenas: Buenos Aires, Rey Curré, Camino a Sabana Mamey, 200–400 m, 16 October 1992, S. Rojas & L.M.
Rojas 102 (F, MO). PANAMA. Colón: Ca. 2–3 miles on Pipeline road, north of Gamboa, 0–10 m, 1 September
1981, S. Knapp 1053 (NY, MO). Darién: Rio Balsa, between Manene and Tusijuanda, 26 July 1967, J.A. Ducke
13544 (3) (MO); 26 July 1967, J.A. Ducke 13579 (3) (MO). El Real, 3 March 1972, A.H. Gentry 4538 (BM, MO).
Rio Tuira between Boca de Cupe and mouth of Rio Pucro, 12 January 1975, A.H. Gentry & S. Mori 13527 (F, MO).
Santa Fe, s.d., J.A. Ducke 8396 (1) (MO). Panamá: Barro Colorado Island, Gigant Bay, 2 August 1934, O.
Shattuck 1108 (F, MO, US); Shore, east side of Barrunga Peninsula, 27 July 1969, R. Foster 1178 (F, MO); Drayton
House Clearing, 23 November 1970, T.B. Cro at 12681 (F photo, MO, NY); North shore of Gigant Bay, 8 March
1971, T.B . Croat 13975 (F, MO, NY); Gigant Bay, 5 April 1971, A.H. Gentry 708 (MO); Vicinity of Laboratory
Clearing, fairchild ridge above boatman's house, 19 October 1973, G. Montgomery 195 (MO); Ibid., near butterfly
cage on east bank of Allee Stream in treefall, 24 October 1973, G. M o n t g o m e r y 1 9 9 (MO). Alhajuela, July 1961,
J.D. Dwyer 1144 (MO). Madden Forest Road at entrance to Boy Scout Road Vine, 27 December 1970, T.B. Cro at
12904 (MO). Boy Scout Camp Road near Madden Lake, 13 October 1971, A.H. Gentry 2058 (F, MO); 12 July
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1972, A.H. Gentry 5502 (F photo, MO); 100 m, 19 December 1972, A.H. Gentry 6696 (MO, NY). Mouth of Rio
Pasiga near coast in cutover forest, 26 October 1971, A.H. Gentry 2203 (F, MO). Rio Pasiga to above waterfall on
second main fork, 29 October 1971, A.H. Gentry 2271 (F, MO). First bend of Rio Pasiga, edges of clearing around
Indian’s house, 1 November 1971, A.H. Gentry 2365 (MO, NY). 0 to 4 km fromo Bayano crossing on road to
Santa Fé, 26 January 1972, A.H. Gentry 3875 (MO). Near archeological site at edge of Madden Lake, 9 April 1972,
A.H. Gentry 5024 (MO). Along stream ca. 3 miles E of Transisthmian highway on road to Salamanca, 100 m, 19
December 1972, A.H. Gentry 6723 (MO). Bordeando el Majé, campamento del G.M.I. isla Bayano, 22 August
1976, C. Garibaldi 225 (MO).
Taxonomic notes:Tynanthus croatianus can be recognized by its ovate to elliptic leaflets and dense
inflorescences. These features are also found in T. densiflorus, but the occurrence of interpetiolar patelliform
glands (versus absent in T. c roa tianus ) differentiate these species. Tyn anthus cro ati anus is morphologically very
distinct from its sister species, T. guatemalensis, another species from Central America (Medeiros & Lohmann
2015). Both species share pubescent to glabrescent leaflets but T. c roat ianus can be easily separated by the trifid
tendril (versus simple in T. guatemalensis), minute prophylls of the axillary buds (versus foliaceous in T.
guatemalensis), corolla 1.2–2 cm long, 4–7 mm wide (versus 0.5–0.9 cm long, 1.9–3.5 mm wide in T.
guatemalensis), and unwinged fruit margins (versus winged in T. guatemalensis).
3. Tynanthus densiflorus M.C. Medeiros & L.G. Lohmann (2014: 79) Type:—BRAZIL. Amazonas: Reserva
Florestal Adolpho Ducke, Manaus–Itacoatiara, km 26, 16 August 1996, L.C. Procópio et al. 14 (holotype INPA!
(189631); isotypes G!, K!, MO! (6223709), NY!, RB! (barcode 58359), SP! (341845)).
Fig. 4: A–I
Lianas. Branchlets subtetragonal to terete, without ritidome, finely striated, lenticeled to densely lenticeled,
pubescent to puberulent, with simple and peltate trichomes; interpetiolar ridge absent or present; interpetiolar
patelliform glands present; prophylls of the axillary buds 0.5–0.8 mm long, 1–2.5 mm wide, minute, shallowly
triangular, puberulent throughout, with simple and peltate trichomes. Leaves 2–3 foliolated; terminal leaflets often
modified into trifid tendrils, without adhesive-disks on tip; petioles and petiolules puberulent to glabrescent
throughout surface, with simple and peltate trichomes; petioles 1.8–5.6 cm long; petiolules (0.6–)1.4–3.8 cm long;
leaflets (3.2–)5–16.1 cm long, (1.3–)2–9.5 cm wide, membranous to chartaceous (sometimes subcoriaceous),
discolor or concolor, ovate; apex caudate, mucronate; base cuneate to truncate or subcordate, symmetrical or
asymmetrical; margin entire; the abaxial surface pubescent to puberulent throughout (sometimes only on and near
the veins), with simple, peltate and patelliform trichomes; the adaxial surface pubescent to glabrescent throughout
(sometimes only on and near the veins), with simple, peltate and patelliform trichomes; glandular trichomes evenly
distributed throughout both surfaces; second venation weak brochidodromous; pocket domatia with (sometimes
without) trichomes. Inflorescence axilar, a thyrse, dense, with corymbose, subcorymbose or conical aspect, 3–9.5
cm long; axis densely pubescent to puberulent, with simple and peltate trichomes; bracts of the inflorescence
predominantly caducous, densely pubescent to pubescent throughout, 0.5–2.5 mm long; floral bracts 0.4–0.6 mm
long; floral pedicels 1–7 mm long. Calyx green to grayish, 1.5–2.2 mm long, 1.4–1.9 mm wide, with transversal
aperture, truncate or minutely 5-denticulate, densely pubescent to pubescent throughout outside, without
patelliform glands; lobes 0.1–0.2 mm long. Corolla cream or pale yellow, 0.8–1.5 cm long, 3–5 mm wide at the
tube opening; tube 3–5 mm long, internally tomentose at the base, with simple and long and short stipitate
trichomes; nectar guides present, yellow; lobes densely pubescent to pubescent throughout lower ones and at
margins of or throughout upper ones; upper ones 0.4–1.4(–2.9) mm long, 0.7–1.5(–2.4) mm wide, acute to obtuse;
lower ones 2.1–4 mm long, 2–3.6 mm wide, obtuse to rounded. Androecium with fertile stamens inserted 1.5–2.5
mm from the base of the corolla; shorter ones 3.5–5.5 mm long; longer ones 4.5–7 mm long; anthers thecae 1.1–1.4
mm long, obovate to elliptic, subexserted; connective extending 0.2–0.3 mm beyond anther attachment; staminode
1.5–2.7 mm long, with long and short stipitate trichomes. Gynoecium 7–9 mm long; ovary 1.3–1.5 mm long,
0.7–0.8 mm wide, conical, velutinous; style 5–7 mm long, tomentose at the base. Fruit not seen. Seeds not seen.
Phenology:—Flowers in August; the fruiting season is unknown.
Distribution and habitat:—Occurs in moist broadleaf forests from Brazil (Amazonas) (Fig. 5).
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TAXONOMIC REVISION OF TYNANTHUS (BIGNONIEAE)
FIGURE 4. Tyna nthu s den siflo rus: A. Flowering branch; B. Detail of lenticels in the oldest portion of branchlet; C. Detail of
pubescent indumentum in the youngest portion of branchlet; D–E. Interpetiolar glands; F. Detail of inflorescence axis with bracts; G.
Open corolla showing the androecium; H. Open calyx showing the gynoecium; I. Ovary cross section showing ovules (L.C. Procópio
14, NY). Illustration by Klei Sousa. Figure from Medeiros & Lohmann (2014).
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FIGURE 5. Distribution of Tynanthus densiflorus.
Additional specimens examined:—BRAZIL. Amazonas: 2–5 km N of Manaus–Itacoatiara Road at km 79
near Rio Preto da Eva, 100–200 m, 24 November 1974, A.H. Gentry 12849 (INPA photo, MG, MO). Rio Camanau,
28 June 1987, P. Grenand et al. 2787 (INPA). Manaus, Campus of INPA, Estrada do Aleixo, 22 November 1974,
A.H. Gentry 12792 (INPA photo); 30 November 1974, A.H. Gentry 13018 (INPA, MO); Ibid., Transect vouchers,
Line 1, 11 December 1974, A.H. Gentry 13181 (INPA, MO); Estrada do Aleixo near Manaus, km 6–7 past INPA, 2
December 1974, A.H. Gentry 13040 (INPA photo, MO); Reserva Florestal Ducke, Parcela PPBio (L03 1000 m),
100 m, 14 December 2010, M.C. Medeiros et al. 21 (SPF); Ibid., próximo à estação meteorológica, 120 m, 15
December 2010, M.C. Medeiros et al. 22 (SPF); Ibid., proximidades do refeitório da base da reserva, na beira da
estrada, 110 m, 16 December 2010, M.C. Medeiros et al. 25 (SPF).
Taxon om ic n ot es: Tynanthus densiflorus is characterized by interpetiolar patelliform glands and dense
inflorescences, with corymbose, subcorymbose or conical aspect (Medeiros & Lohmann 2014). Tynanthus
densiflorus is closely related to T. panurensis and T. pubes cens (Medeiros & Lohmann 2015). It shares ovate
leaflets and an internally tomentose corolla tube at the base with T. panu rensis; however, T. densiflor us is easily
identified by the presence of interpetiolar glands (versus absent in T. panurensis), minute prophylls of the axillary
buds (versus foliaceous in T. panurensis ) and dense inflorescences (versus lax in T. panurensis) (Medeiros &
Lohmann 2014). On the other hand, T. densif lorus shares interpetiolar patelliform glands and similar corolla length
with T. pubescens (i.e., 1–1.6 cm in T. pubescens and 0.8–1.5 cm in T. densiflorus), but differs by the caudate-
mucronate leaflet apices (versus acuminate or obtuse-mucronate in T. pubescens) and dense inflorescences (versus
lax in T. pubescens) (Medeiros & Lohmann 2014).
4. Tynanthus espiritosantensis M.C. Medeiros & L.G. Lohmann (2014: 82) Type:—BRAZIL. Espírito Santo:
Linhares, Reserva Natural da CVRD, Estrada Oiticica, km 2.3, 6 February 2008, D.A. Folli 5931 (holotype CVRD!
(11073); isotype SPF! (barcode 199170)).
Fig. 6: A–I
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TAXONOMIC REVISION OF TYNANTHUS (BIGNONIEAE)
FIGURE 6. Tyn ant hus e spi ritos ant ensi s: A. Flowering branch; B–D. Leaflet with pubescent domatia in the abaxial surface; E.
Interpetiolar region with bromeliad-like prophylls of the axillary buds; F. Detail of the inflorescence axis, showing bracts, simple and
peltate trichomes; G. Open corolla showing the androecium; H. Anther; I. Open calyx showing the gynoecium (D.A. Folli 5931, SPF).
Illustration by Klei Sousa. Figure from Medeiros & Lohmann (2014).
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Lianas. Branchlets tetragonal to terete, with or without ritidome, finely striated, lenticeled to densely lenticeled,
glabrescent (sometimes pubescent at the nodes), with peltate and patelliform trichomes (sometimes also with
simple); interpetiolar ridge absent; interpetiolar patelliform glands absent; prophylls of the axillary buds 1.2–2.5
mm long, 0.7–1.1 mm wide, bromeliad-like, glabrescent (rarely puberulent) throughout, with peltate trichomes
(rarely also with simple). Leaves 2–3 foliolated (more commonly 3); terminal leaflets often modified into trifid
tendrils, without adhesive-disks on tip; petioles and petiolules puberulent throughout surface, with simple and
peltate trichomes; petioles 1–6 cm long; petiolules 0.5–3.5 cm long; leaflets (4–)5–11.9 cm long, (1.5–)1.9–5.4 cm
wide, membranous to chartaceous, discolor, elliptic; apex acuminate or caudate, mucronate; base cuneate,
symmetrical; margin entire; the abaxial surface glabrescent (sometimes pubescent) on and near the veins, with
peltate and patelliform trichomes (sometimes also simple); the adaxial surface glabrescent on and near the veins,
with peltate and patelliform trichomes; glandular trichomes distributed especially on the abaxial surface; second
venation weak brochidodromous; pocket domatia with trichomes. Inflorescence axilar, a thyrse, lax, with conical
aspect, 3.6–7 cm long; axis pubescent, with simple and peltate trichomes; bracts of the inflorescence predominantly
caducous, pubescent throughout or only at margins, 0.7–3.9(–9) mm long; floral bracts 0.5–0.7 mm long; floral
pedicels 3.5–9 mm long. Calyx green, 2.3–2.7 mm long, 1.8–2.5 mm wide, with transversal (sometimes oblique)
aperture, minutely 5-denticulate, glabrescent (sometimes pubescent at teeth) outside, with patelliform glands; lobes
0.1–0.4 mm long. Corolla white, 0.7–0.8 cm long, 2.5–3.4 mm wide at the tube opening; tube 2.5–4 mm long,
internally tomentose to pubescent at the base or glabrescent, with simple and long and short stipitate trichomes;
nectar guides absent, but with a path of long and short stipitate trichomes; lobes densely pubescent to pubescent
throughout lower ones and at the margin of upper ones; upper ones 0.4–1.1 mm long, 1–1.5 mm wide, acute to
obtuse; lower ones 1.8–3.2 mm long, 2.1–2.5 mm wide, obtuse to rounded (sometimes acute). Androecium with
fertile stamens inserted 1–1.5 mm from the base of the corolla; shorter ones 2.5–3.5 mm long; longer ones 4.5–5
mm long; anthers thecae 0.8–1.1 mm long, obovate to elliptic, subexserted; connective extending 0.2–0.3 mm
beyond anther attachment; staminode ca. 2.4 mm long, glabrescent, with long and short stipitate trichomes.
Gynoecium 4.5–6 mm long; ovary 0.8–1 mm long, 0.7–0.9 mm wide, conical, velutinous; style 3.3–5 mm long,
tomentose at the base. Fruit not seen. Seeds not seen.
Phenology:—Flowers from December to February; the fruiting season is unknown.
Distribution and habitat:—Occurs in moist broadleaf forests from Brazil (Espírito Santo) (Fig. 7).
FIGURE 7. Distribution of Tyn ant hu s es pir it os an ten si s.
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TAXONOMIC REVISION OF TYNANTHUS (BIGNONIEAE)
Additional specimens examined:—BRAZIL. Espírito Santo: Linhares, Rancho Alto, 7 December 1984, G.
Hatschbach & J.M. Silva 48693 (MBM, MO, US); Reserva Natural da CVRD, Estrada Oiticica, próximo à
porteira, antes do cruzamento com a estrada municipal, 53 m, 27 January 2014, M.C. Medeiros & R.B. Louzada 41
(CVRD, SPF).
Taxo n om ic no te s :Tynanthus espiritosantensis shares bromeliad-like prophylls of the axillary buds,
glabrescent elliptic leaflets, corolla tube internally tomentose at base and lax inflorescences with T.
schumannianus. Nevertheless, T. espir itosant ensis is easily separated by the pubescent leaflet domatia (versus
glabrous domatia in T. schumannianus), lack of patelliform glands in the petioles, petiolules and inflorescence axis
(versus presence of patteliform glands in T. schumannianus) and larger calyx, with 2.3–2.7 × 1.8–2.5 mm (versus
smaller calyx, with 1–2 × 1.1–1.9 mm in T. schumannianus) (Medeiros & Lohmann 2014). Additionally, T.
espiritosantensis occurs in the Atlantic Forest of Espírito Santo, while T. schumannianus is an Amazonian species.
Apart from their morphological similarity, T. espiritosant ensis and T. schuma nnianus are also closely related,
belonging to a clade that also contains T. fasciculatus, T. labiatus and T. micranthus , all distributed through the
Atlantic Forest (Medeiros & Lohmann 2015).
5. Tynanthus fasciculatus (Vell.) Miers (1863: 193). Bignonia fasciculata Vellozo (1825: 247; 1827: tab. 25).
Arrabidaea fasciculata (Vell.) de Candolle (1845: 185). Cuspidaria fasciculata (Vell.) Sonder (1849: 560).
Schizopsis fasciculata (Vell.) Bureau (1865: 379). Lectotype (designated by Lohmann, in Lohmann & Taylor 2014:
468):—Fl. Flumin. Icones 6: tab. 25. 1827.
Fig. 8: A–F
Lianas. Branchlets tetragonal to terete, with or without ritidome, finely striated, densely lenticeled, tomentose to
pubescent, with simple and peltate trichomes; interpetiolar ridge absent or present; interpetiolar patelliform glands
absent; prophylls of the axillary buds 0.7–1.5(–2.2) mm long, 0.5–1.2(–1.5) mm wide, minute, shallowly triangular
to triangular, tomentose to pubescent throughout, with simple and peltate trichomes. Leaves 2–3 foliolated (more
commonly 3); terminal leaflets often modified into trifid tendrils, without adhesive-disks on tip; petioles and
petiolules tomentose to pubescent throughout surface, with simple and peltate trichomes; petioles (0.5–)1.2–5.5 cm
long; petiolules (0.4–)0.6–3.4 cm long; leaflets (3.3–)4.3–11.7 cm long, (1.2–)2–7.5 cm wide, membranous to
chartaceous (sometimes subcoriaceous), discolor or concolor, elliptic; apex acuminate, mucronate; base cuneate or
obtuse, symmetrical; margin entire; the abaxial surface tomentose to pubescent throughout or only on and near the
veins, with simple and peltate trichomes; the adaxial surface tomentose to pubescent throughout or only on and
near the veins, with simple and peltate trichomes; glandular trichomes evenly distributed throughout both surfaces;
second venation weak brochidodromous; pocket domatia with trichomes. Inflorescence axilar or terminal, a thyrse,
dense, with corymbose or subcorymbose aspect, (2–)3.1–4.5 cm long; axis tomentose, with simple and peltate
trichomes; bracts of the inflorescence predominantly caducous, tomentose throughout, 1–3.5 mm long; floral bracts
0.6–1.4 mm long; floral pedicels 1.5–6 mm long. Calyx green to reddish, 2.5–4 mm long, 2–3.5 mm wide, with
transversal (sometimes oblique) aperture, 5-laciniate, tomentose or densely pubescent throughout outside, with
patelliform glands; lobes 0.6–3 mm long. Corolla white, cream or pale yellow, 1.2–2.2 cm long, 4–7 mm wide at
the tube opening; tube 5–11 mm long, internally tomentose to pubescent at the base, with simple and long and short
stipitate trichomes; nectar guides present, yellow; lobes densely pubescent to pubescent throughout lower ones and
at the margin of upper ones; upper ones 0.7–2.8 mm long, 1–3.3 mm wide, acute to obtuse; lower ones 4–5.2 mm
long, 2.5–5.2 mm wide, acute, obtuse or rounded. Androecium with fertile stamens inserted 2.5–3 mm from the
base of the corolla; shorter ones 7.5–8.5 mm long; longer ones 9–10 mm long; anthers thecae 1.4–1.7 mm long,
obovate to elliptic, subexserted; connective extending 0.4–0.6 mm beyond anther attachment; staminode 2.3–2.8
mm long, glabrescent, with long and short stipitate trichomes. Gynoecium 12–13 mm long; ovary 1.5–2 mm long,
0.9–1.3 mm wide, conical, velutinous; style 10–12 mm long, tomentose at the base. Fruit a linear flattened to
subtetragonal capsule, 11–39.5 cm long, (0.5–)0.8–2 cm wide, woody, smooth to granular near the midvein and
granular near the margins, without lenticels to densely lenticeled, tomentose to pubescent, with simple, peltate and
patelliform trichomes; central ridge single, prominent (sometimes not); margins prominently raised (winged),
0.2–0.7 cm wide. Seeds body (0.7–)1–2 cm long, (0.4–)0.6–1 cm wide; wings (0.3–)0.5–1.2 cm long.
Phenology:—Flowers from September to December and produces fruits from February to October.
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FIGURE 8. A–F. Tyn an thus f as cic ul at us : A. Flowering branch; B. Detail of tomentose indumentum in branchlets; C. Calyx with
laciniate apex; D. Open corolla showing the androecium; E. Gynoecium (G. Edwall s.n., F 896072); F. Fruit showing the subtetragonal
cross section (M. Barreto 1824, F). G–J. T. guat em al en sis: G. Flowering branch; H. Interpetiolar region showing the foliaceous
prophylls of the axillary buds; I. Flower (E. Contreras 8517, F); J. Fruit (P.H. Gentle 7775, F). Illustration by Klei Sousa.
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TAXONOMIC REVISION OF TYNANTHUS (BIGNONIEAE)
Distribution and habitats:—Occurs mainly in moist broadleaf forests, although a few populations are known
to occur in savanna areas from Eastern Brazil (Minas Gerais and São Paulo) (Fig. 9).
FIGURE 9. Distribution of Tyn ant hu s fa sci cu la tus.
Additional specimens examined:—BRAZIL. Sin loc., s.d., Araujo? 11364 (P barcode 3606710). Sin loc.,
s.d., J.S. Blanchet s.n. (BM). Sin loc., s.d., J.C. Mello s.n. (NY barcode 483766, US 2515368). Sin loc., s.d.,
Widgren 243 (BM). Sin loc., 1 September 1866, J.C. Mello 8a (K, P barcode 3606703). Sin loc., 18 October 1866,
J.C. Mello 8β (P barcode 3606704). Sin loc., 1 October 1867, J.C. Mello 8y (K, MO 3395613, P barcode 3606705,
US 2515615). Minas Gerais: Sin loc., s.d., A.F. Regnell II-198 (MO 3395612). Sin loc., 1845, Widgren s.n. (BR,
MO 2698894, MO 3395608, P barcode 3606700). Jardim, 20 November 1845, Widgren 49 (BR). Sin loc., 1845,
Widgren 94 (BR). Sin loc., 18 October 1847, A.F. Regnell II-198 (K, LE). Sin loc., 19 October 1861, A.F. Regnell
II-198 (R 23787, US 201375). Rio Pardo, 7 July 1866, A.F. Regnell II-198 (K). Sin loc., 9 September 1866, A.F.
Regnell II-198 (P barcode 3606695). Serra da Concha, 21 September 1867, A.F. Regnell II-198 (P barcode
3606699); 29 September 1867, A.F. Regnell II-198 (P barcode 3606698). Campos da Bocaina, 8 September 1879,
A. Glaziou 4692a (P, US). Perpetua, pres Diamantina, 11 April 1892, A. Glaziou 19664 (P). Mineração da Itaú,
Rodovia Belo Horizonte a Paraopeba, 13 September 1957, E.P. Heringer s.n. (RB barcode 58281; RB barcode
58305; MO 2609963). Belo Horizonte, Estação Ecológica UFMG, Trilha J, 6 June 1990, E.M. Santos et al. s.n. (F
2170556); 6 June 1990, E.M. Santos et al. 76 (BHCB); Ibid., 2a. Estação de coleta, trilha J, 10 October 1990, E.
Tameirão Neto 215 (BHCB, F). Caldas, s.d., A.F. Regnell II-198 (K, MO 2305639, P barcode 3606697, R 23786,
US 2515367); 8 September 1845, A.F. Regnell II-198 (P barcode 3606701); 19 October 1861, A.F. Regnell II-198
(P barcode 3606694); 21 September 1867, A.F. Regnell II-198 (BR, M); 1869, A.F. Regnell II-198 (BR).
Camanducaia, Mata dos Mota, 1000 m, 12 October 1999, R.B. Torres et al. 756 (IAC). Coronel Pacheco, s.d., E.P.
Heringer s.n. (RB barcode 58310); Estação Experimental, 28 October 1940, E.P. Heringer 352 (RB, SP, VIC). Juiz
de Fora, Serra do 12º RI, 11 November 1964, S.V. Monteiro 2627 (VIC). Marliérea, Parque Estadual do Rio Doce,
Salão Dourado, junto à saída, 15 June 1995, J.A. Lombardi & L.G. Temponi 793 (BHCB). Mercês, 1840?, G.
Gardner 5030 (BM, K, G, P). Ouro Preto, Parque Estadual do Itacolomi, Trilha da Mata do sibrão, 11 December
2006, R.S. Araújo et al. 44 (VIC). Reduto, Reduto E.F.L., 12 October 1950, E.P. Heringer 2656 (NY, RB). Tombos,
Fazenda da Cachoeira, 9 August 1935, M. Barreto 1824 (F, MO, R). Uberaba, 7 July 1860, A.F. Regnell II-198 (US
1322404). Viçosa, Direction of São Miguel, km 8, Fazenda A. Cocerro, 900 m, 26 June 1930, Y. Me xi a 4 80 2 (BM,
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F, G, K, MO, NY, P, US); Mata do Paraíso, 5 December 1935, J.G. Kuhlmann s.n. (RB barcode 58468; VIC 2672);
Ibid., aceiro, em baixada ca. de 100 m depois da segunda cerca, subindo à esquerda da cancela, 23 October 2002,
M.M.M. Lopes 158 (VIC); 17 February 2003, M.M.M. Lopes 174 (VIC); UFV, 23 February 1959, H.S. Irwin 2714
(F photo, K, US); Ibid., Jardim Botânico, 7 October 1962, M.R.R. Vidal 199 (VIC); 15 October 1979, A.J. Filho s.n.
(VIC 6393); 15 October 1979, R.S. Ramalho 1621 (RB); 11 November 2010, M.C. Medeiros & R.B. Louzada 19
(SPF). São Paulo: Sin loc., September 1868, S. Henschen s.n. (US 201374). Estrada de Pirapora a Cabreúva, 4
December 1924, A. Gehrt s.n. (NY barcode 483765, SP 12903, US 1543120). Caieiras, Terrenos da Cia.
Melhoramentos de São Paulo, 27 October 1936, M. Kuhlmann s.n. (SP 36635). Campinas, Souzas, s.d., H.M.
Souza s.n. (IAC 20158); 4 May 1866, J.C. Mello 8 (K, P barcode 3606702); 20 July 1875, H. Mosén 3960 (MO, P);
Barão Geraldo, Santa Genebra Forest Reserve, Betel, Transect 1, 600 m, 4 January 1985, A.H. Gentry & E. Zardini
49163 (MO); 4 January 1985, A.H. Gentry & E. Zardini 49173 (MO); Ibid., Transect 2, 550 m, 26 August 1987,
A.H. Gentry & A. Silva 58692 (MO, UEC); Ibid., 550 m, 27 August 1987, A.H. Gentry & A. Silva 58713 (MO,
UEC); Ibid., Transect 5, 550 m, 31 August 1987, A.H. Gentry 58780 (MO, UEC); Ibid., 550 m, 5 September 1987,
A.H. Gentry 59067 (K, MO, NY, UEC); Ibid., 550 m, 5 September 1987, A.H. Gentry 59077 (UEC); Sítio São
Francisco, 31 January 2000, T. Spinelli et al. 219 (UEC). Helvetia, 3 November 1943, D.B.J. Pickel s.n. (SPF
barcode 200750, SPSF 948, US 1564384); Ibid., 10 November 1943, D.I. Stehle s.n. (SPSF 1012). Jundiai, E.E.
Jundiaí, 12 April 1994, L.C. Bernacci et al. 1 (HRCB, SPF). Limeira, Orla da mata da S.A.F.B, 10 October 1946,
M. Kuhlmann 1278 (US). Rio Claro, Fazenda São José, 24 August 2000, M.A. Assis et al. 1367 (HRCB); Ibid., 4
October 2000, R. Udulutsch & V.T. Rampin 57 (HRCB); Ibid., 15 November 2000, R. Udulutsch & M.A. Assis 112
(HRCB); Ibid., 11 January 2001, R. Udulutsch & M.A. Assis 169 (HRCB); Ibid., 24 March 2001, R. Udulutsch et
al. 232 (HRCB); Ibid., 24 March 2001, R. Udulutsch et al. 245 (HRCB); Ibid., 7 October 2001, M.A. Assis & A.G.
Manzatto 1595 (HRCB); Ibid., 30 October 2001, R. Udulutsch et al. 429 (HRCB); Ibid., 3 October 2003, M.A.
Assis et al. 1666 (HRCB). Santa Rita do Passa Quatro, Parque Estadual de Vassununga, Gleba Maravilha, 17
December 2002, Y. J. A. Ti bi ri çá & L .F. Co el ho 70 (HRCB). São Paulo, Cidade Jardim, 13 November 1941, W.
Hoehne 804 (G, RB). Serra Negra, October–November 1901, G. Edwall s.n. (F 896072, SP 15045).
Taxonomic notes:Tyna nthus f asciculatus can be easily recognized by the laciniate calyx that contrasts with
the denticulate or truncate calyx apices found in the remaining species of the genus. This species is the only
Southern Brazilian species to bear dense inflorescences, with a corymbose or subcorymbose aspect. Tynanthus
fasciculatus is sister to T. labiatus, another Atlantic Forest species (Medeiros & Lohmann 2015), with which it
shares winged fruits. However, T. fasciculat us can be easily separated by the dense inflorescences (versus lax
inflorescences in T. labiatus) and the dense indumentum on young branchlets (versus glabrescent branchlets in T.
labiatus).
6. Tynanthus guatemalensis Smith (1893: 6). Lectotype (designated here):—GUATEMALA. Quezaltenango: Banks
of Rio Ocosito, April 1892, J. Donnell Smith 1488 (US! (barcode 125785); isolectotypes GH! (barcode 93265) photo,
K! (barcode 449544), M!, NY! (barcode 328979), P! (barcode 468599), US! (barcode 125786)).
Fig. 8: G–J
Lianas. Branchlets tetragonal to terete, with or without ritidome, finely striated, lenticeled to densely lenticeled,
pubescent to glabrescent, with simple and peltate trichomes; interpetiolar ridge present; interpetiolar patelliform
glands absent; prophylls of the axillary buds 3.5–20 mm long, 2.1–14 mm wide, foliaceous, ovate to elliptic,
pubescent or puberulent to glabrescent throughout, with simple, peltate and patelliform trichomes. Leaves 2–3
foliolated (more commonly 3); terminal leaflets often modified into simple tendrils, without adhesive-disks on tip;
petioles and petiolules pubescent throughout surface, with simple and peltate trichomes; petioles (0.9–)1.5–5.6 cm
long; petiolules (0.4–)0.6–3.4 cm long; leaflets (2.5–)3.5–12.2 cm long, (1.2–)1.9–7.9 cm wide, membranous to
chartaceous (sometimes subcoriaceous), discolor or concolor, elliptic; apex caudate, mucronate; base cuneate or
obtuse, symmetrical; margin entire; the abaxial surface pubescent to glabrescent on and near the veins, with simple,
peltate and patelliform trichomes; the adaxial surface pubescent to glabrescent on and near the veins, with simple,
peltate and patelliform trichomes; glandular trichomes distributed especially on the abaxial surface; second
venation weak brochidodromous; pocket domatia without (sometimes with) trichomes. Inflorescence axilar or
terminal, a thyrse or a compound thyrse, lax, with conical aspect, first order (1.5–)2.6–10.5 cm long, second order
0.5–2.1 cm long; axis pubescent, with simple and peltate (sometimes also with patelliform) trichomes; bracts of the
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TAXONOMIC REVISION OF TYNANTHUS (BIGNONIEAE)
inflorescence caducous or persistent, pubescent throughout (sometimes only at margins), 0.4–3.3 mm long; floral
bracts 0.3–0.5 mm long; floral pedicels 1–4 mm long. Calyx green, 1.5–2.5 mm long, 1.3–2.5 mm wide, with
transversal aperture, truncate or minutely 5-denticulate, pubescent to puberulent (sometimes glabrescent)
throughout outside, with patelliform glands; lobes 0.1–0.4(–0.5) mm long. Corolla white, cream or pale yellow,
0.5–0.9 cm long, 1.9–3.5 mm wide at the tube opening; tube 2–5 mm long, internally pubescent at the base, with
simple and long and short stipitate trichomes; nectar guides present, yellow; lobes densely pubescent to pubescent
throughout lower ones and at the margin of (sometimes throughout) upper ones; upper ones 0.3–1.8 mm long,
0.6–1.9 mm wide, acute to obtuse; lower ones 1.4–3.1 mm long, 1.6–3.2 mm wide, obtuse to rounded. Androecium
with fertile stamens inserted ca. 1 mm from the base of the corolla; shorter ones 2–4 mm long; longer ones 3–5 mm
long; anthers thecae 0.6–1 mm long, obovate to elliptic, subexserted; connective extending 0.1–0.3 mm beyond
anther attachment; staminode 1.7–2 mm long, glabrous. Gynoecium 4–6.5 mm long; ovary 1–1.4 mm long, 0.6–0.8
mm wide, conical, densely pubescent; style 3–5 mm long, tomentose to pubescent at the base. Fruit a linear
flattened capsule, (12.7–)14–38 cm long, 0.8–2.4 cm wide, woody, smooth or granular throughout, without
lenticels to densely lenticeled, glabrescent (sometimes pubescent at extremities), with peltate and patelliform
trichomes (sometimes also with simple); central ridge single, slightly or not prominent; margins prominently raised
(winged), 0.3–0.9 cm wide. Seeds body 0.8–2 cm long, 0.5–0.9 cm wide; wings 0.6–1.8 cm long.
Phenology:—Flowers from April to July and produces fruits from August to April.
Distribution and habitat:—Occurs in moist broadleaf forests from Belize (Belize, Cayo, Orange Walk, Stann
Creek and Toledo), Guatemala (Alta Verapaz, El Petén, Izabal and Quetzaltenango) and Mexico (Campeche,
Chiapas, Quintana Roo, Tabasco and Veracruz-Llave) (Fig. 10).
FIGURE 10. Distribution of Tyna nth us gua tem ale nsi s.
Additional specimens examined:—BELIZE. Belize: Gracy Rock Bank (“Gracie Rock”), Sibun River, 27
May 1935, P.H. Gentle 1652 (K, MO, NY, P). Maskall, 25 May 1934, P.H. Gentle 1324 (F, K, MO, NY, US). South
of Yalbac Hills, Terra Nova Medicinal Plant Reserve, 40 m, 11 June 1993, J.B. Walker & L. Romero 138 (NY).
Cayo: Cayo village, May 1907, E. Campbell 112 (K). Arenal–Valentin road, June–August 1936, C.L. Lundell 6168
(F, NY, US). 53 Miles Section, Humming Bird Highway, 18 June 1956, P.H. Gentle 9132 (F, K, MO). Maccal
River, bridge between Augustine and Cuevas, 7 October 1972, J.D. Dwyer & R. Pippin 10189 (MO). Vicinity of
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Cuevas south of Millionario, 29–30 May 1973, T.B . Croa t 23599 (F, MO, NY). South of Millionario, 1900 ft, 29
May 1973, A.H. Gentry 7661 (F, INPA, MO, NY, US). Vicinity of Millionario, 1800 ft, 30 May 1973, A.H. Gentry
7689 (MO, NY); 30 May 1973, A.H. Gentry 7730 (F, MO, NY). Between Cuevas & Millionario, 30 May 1973,
J.D. Dwyer 10814 (MO, US). Vicinity of Millionario between the McCal River and Cuevas, 1900 ft, 30 May–3
June 1973, T.B. Croat 23715 (F, K, MO, NY, US). 3 miles S of Grano de Oro, 1700 ft, 2 June 1973, A.H. Gentry
7755 (F, MO, NY). Vicinity of Grano de Oro lumber camp south of Millionario, 1700 ft, 2 June 1973, A.H. Gentry
7768 (F, MO, NY). 4 miles south of Grano de Oro on road to La Flor, 2 June 1973, J.D. Dwyer 10923 (MO, US).
Vicinity of La Flor at Rio de La Flor, 6 mi. south of Grano de Oro, 1700–2000 ft, 3 June 1973, T.B. Cro at 2 381 2 (F,
MO). Western Highway, near Belmopan, 3 June 1981, C. Whitefoord 3115 (BM, MO). Monkey Tail track, 450 m,
29 May 1995, C. Whitefoord 9499 (BM, F, MO). Chiquibul Forest Reserve, San Pastor track, c. 1 km from Las
Cuevas, 560 m, 10 June 1996, A.I. Garcia A68 (BM, MO). Augustine, Rio Frio Caves road, Mountain Pine Ridge
Forest Reserve, 460 m, 3 July 1989, J. Meave & A. Howe 1433 (BM, MO). Valentin, June–July 1936, C.L. Lundell
6354 (F, NY, US). Orange Walk: Mile 58 on Northern Highway, ca. 4 miles south of Tower Hill, 23 June 1973,
A.H. Gentry 8489 (MO). 1.5 km S of Program for Belize Camp, 13 May 1991, R. Arvigo et al. 502 (NY).
Programme for Belize Rio Bravo Research Station, 150 m, 28 May 1997, D. Lentz et al. 2744 (NY). Stann Creek:
Stann Creek Valley, Mountain Cow Ridge, 25 March 1940, P.H. Gentle 3275 (MO, NY, US). Cockscomb
Mountains, tributary of Cocoa Branch of Sittee River, 2 km due north of Victoria Peak, 300–500 ft, 5–6 June 1973,
A.H. Gentry 8014 (MO). Toledo: Temash River, 200 ft, 7 May 1935, W.A . Sch ipp 8910 (F photo); 5 June 1944,
P.H. Gentle 4654 (MO, NY). Near Agua Caliente, near San Antonio, 18 April 1945, P.H. Gentle 5335 (F, K, MO).
Beyond Central Camp, Edwards Road beyond Columbia, 21 May 1951, P.H. Gentle 7339 (F, K, MO, NY). Broken
ridge, Joe Taylor creek, 2 October 1952, P.H. Gentle 7775 (F, K, MO). 1.5 miles from Maya village of San Jose on
road to Columbia Forestry Station, 12 June 1973, A.H. Gentry 8119 (MO). Mile 13 on road west from Punta Gorda,
near junction of Southern Highway and road to San Antonio, 100 ft, 14 June 1973, A.H. Gentry 8222 (MO).
Southwestern Maya Mountains, Columbia River Forest Reserve, Trail between Gloria Camp and Edwards Camp to
the South, 15 April 1992, B.K. Holst 4468 (MO). GUATEMALA. Alta Verapaz: Along Río Icvolay between Río
Apia and Río Soctelá, 8–10 miles northwest of Cubilgüitz, 200–210 m, 14 March 1942, J.A. Steyermark 45058 (F).
Panzós, Finca Mercedes, Teleman, 70 m, 3 September 1988, E. Martínez S. et al. 23472 (MO). El Petén:
Fallabón–Yaxha Road, 22 March 1933, C.L. Lundell 2193 (F). Carmelita, 1 m south of village, on trail to Flores, 24
June 1942, F. E . E g l e r 4 2 - 2 2 7 (F). Tikal National Park, Tikal, in ramonal on Remate road, 19 May 1959, C.L.
Lundell 15999 (US); Ibid., in zapotal on Remate road, 3 July 1959, C.L. Lundell 16142 (F, G, NY, US); Ibid., in
ramonal covering the ruines, 27 August 1960, E. Contreras 1467 (F, K, MO, NY). Remate, on Tikal Road, in
zapotal, about 12 km NE of the village, 14 May 1960, E. Contreras 942 (US). Dos lagunas, Ixcanrio, on Aguas
Turbias Road, 12 km E, in zapotal, 11 May 1969, E. Contreras 8517 (F). Laguna Yaxja, Banks of Laguna Yaxja
and dry wooded hills of limestone to north and south, 160–250 m, 28 June 1970, W.E . Ha rm on & J .D . Dw ye r 2772
(MO). Dolores, Rio Mopan, 21 February 1971, E. Contreras 10583 (MO, NY). Santa Elena, en la orilla del camino
para San Andrés, a km 17, lado noreste, 25 May 1971, R.T. Ortíz 1782 (F, MO, NY, US). Ca. 5 miles S of Tikal,
logging road 0–4 miles W of road to Tikal, 19 June 1973, A.H. Gentry 8346 (MO, US). Izabal: Trail from Los
Amates to Izabal, 31 May 1919, S.F. Blake 7802 (US). Vicinity of Quiriguá, 75–225 m, 15–31 May 1922, P. C .
Standley 24535 (MO, NY, US). MEXICO. Yucatán, September 1935, R.S. Flores s.n. (F 782747). Campeche:
Calakmul, A 500 m al NW de Dos Naciones, 190 m, 23 October 1997, E. Martínez S. et al. 29324 (NY); A 5 km al
E de La Mancolona, 200 m, 17 November 1997, E. Martínez S. et al. 29720 (NY); 6.5 km al O de Flores Magón,
153 m, 12 March 2002, J.C. Soto et al. 22687 (MBM); A 6.7 km al E de La Nueva Vida, 302 m, 13 May 2003, D.
Álvarez 5026 (MO, SPF); A 4.3 km al O del poblado Flores Magón, 182 m, 3 June 2003, D. Álvarez 5219 (MBM,
MO); A 7.6 km al E del poblado "Unión 20 de Junio" (antes la Mancolona), 160 m, 2 August 2003, D. Álvarez & C.
Jiménez J. 6140 (MO). Hopelchén, A 4 km al O del ejido Santa Rosa, sobre el camino al ejido Carlos A. Madrazo,
100 m, 25 September 1996, P. A l v a r o M . 6 0 5 (BM, MO); A 2.7 km al S de Chan Chen, 12 June 2004, D. Álvarez et
al. 8957 (MO); A 6.64 km al S de Xmejia, 164 m, 15 June 2004, D. Álvarez et al. 9199 (MBM, MO). Chiapas:
Ocosingo, Ruins of Bonampak, 14 March 1975, W. S. Ho ove r 244 (MO, US); En el viejo poblado de Jalisco (Selva
Lacandona), 350 m, 1 December 1976, J.I. Calzada et al. 2907 (MO); South of Santo Domingo on road to
Bonampak and Echeverria from Chancala, 455 m, 25 January 1982, D.E. Breedlove & F. Almeida 57938 (MO);
Siria, a 62 Km al SE de Palenque, 400 m, 12 May 1982, M. Sousa et al. 12397 (BM, MO); A 9 km al NW de Boca
Lacantum, camino a Crucero Corozal–Palenque, 160 m, 16 October 1984, E. Martínez S. 8301 (MO); A 5 km al
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TAXONOMIC REVISION OF TYNANTHUS (BIGNONIEAE)
NO de Crucero Corozal, 170 m, 14 June 1985, E. Martínez S. & G. Aguilar 15000bis (MO); Camino a las ruinas de
Bonampak, reserva Montes Azules, 10 km NE del poblado de Lacanjá, 200 m, 26 October 1985, S.S. Colín & G.I.
Manriquez 319 (MO); En Ejido Roberto Bárrios a 60 km al S de Boca Lacantum, camino a Chajul, 200 m, 18 April
1986, E. Martínez S. 18357 (F, MO); A 15 km al NW de Boca Lacantum, camino a Palenque, 220 m, 12 June 1986,
E. Martínez S. & M.A. Soto A. 18582 (F photo, MO); En Nuevo Chihuahua a 70 km al S de Boca Lacantum, 200 m,
23 June 1986, E. Martínez S. 18966 (F, MO); Borde del río Lacantún, Estación Biológica de Chajul, 150 m, 2
August 1996, G.I. Marínquez et al. 4035 (MO); Nuevo Guerrero, 200 m, 11 April 2002, G. Aguilar M. 472 (MBM,
MO); A 0.2 km de Nuevo Jerusalem camino a Nuevo Francisco León, 12 m, 4 May 2002, G. A g u i l a r M . e t a l . 6 3 0
(MBM, MO); 4 May 2002, G. A g u i l a r M . et a l . 6 8 5 (MO); A 6.5 km al S de Nuevo Guerrero sobre el camino a
Santo Domingo, 400 m, 6 May 2002, D. Álvarez et al. 961 (M, RB); 0.5 km al S de Nuevo Francisco León
(Restaurante El Paraíso), 177 m, 10 May 2002, J.C. Soto et al. 23490 (M, MO); A 1 km al S de El Paraíso, 325 m,
22 May 2002, G. A g u i l a r M . 9 9 2 (MO); A 600 m al SO de Nuevo Jerusalém, 340 m, 31 May 2002, G. A g u i l a r M . e t
al. 1286 (MO); A 200 m de Nuevo Francisco León, restaurante El Paraíso, 174 m, 16 June 2002, G. A g u i l a r M . e t
al. 1399 (M, MBM, MO); A 2 km al N de Nuevo Guerrero, 190 m, 22 June 2002, G. A g u i la r M . e t a l . 1 56 1 (M,
MO, NY); A 1 km de Nuevo México, al Norte del Poblado, 176 m, 9 July 2002, G. A g u i l a r M . & F. A g u i l a r 1 8 0 5
(M, MO); A 3.8 km al SE del paraíso, 385 m, 14 October 2002, G. A g u i l a r M . & D . Á l v a re z 3 4 4 9 (MBM, MO,
NY); A 4 km al S de San Javier, 350 m, 26 January 2003, G. A g u i l a r M . e t a l . 5 2 0 6 (MBM, MO, NY, SPF); A 2.1
km al SE de la comunidad Lacanjá Chansayab, 320 m, 17 June 2003, D. Álvarez 5378 (MO); A 1 km de al NO del
poblado "El limonar", 521 m, 30 August 2003, D. Álvarez & A. Chambor 6297 (MO); A 0.57 km al SO del crucero
de Bonampak, 340 m, 15 October 2003, G. Aguilar M. et al. 8178 (MO); A 0.64 km al O del crucero de Bonampak,
313 m, 13 December 2003, G. Aguilar M. et al. 8908 (MBM). Palenque, Ruinas de Palenque, 160 m, 11 May 1982,
M. Sousa et al. 12382 (BM, MBM, MO). Quintana Roo: A 15 km al Sur de Ejido Laguna OM, 8 June 1980, O.
Téllez & E. Cabrera 2423 (BM, MO, NY). A 8 km al Norte de La Unión, 9 June 1980, O. Téllez & E. Cabrera
2456 (MO, NY). A 12 km al Sur del Ejido Laguna OM, sobre el camino a Tomas Garrido, 7 August 1980, O. Téllez
& E. Cabrera 3054 (MO, NY). A 7 km al norte de Bacalar, 21 December 1982, E. Cabrera & H. de Cabrera 4139
(MO). 4 km al NO de Estero Franco, sobre el camino en construccion a Tomas Garrido, lugar llamado El Danto, 17
May 1985, E. Cabrera et al. 8349 (BM, MO). Adolfo de la Huerta, A 4.2 km al ESE de San Carlos, camino a Pozo
Pirata, 102 m, 20 June 2004, D. Álvarez et al. 9517 (MBM, MO). Othón P. Blanco, A 1.5 km al N de Dos Aguadas,
66 m, 21 June 2004, D. Álvarez et al. 9560 (MO). A 1.8 km al O de El Martirio, camino a Pozo Pirata, 97 m, 9
September 2004, D. Álvarez et al. 10585 (BM). Tabasc o : Balancán, La Palma, 1–6 June 1939, E. Matuda 3277 (F,
K, MO, NY). Huimanguillo, El 32 rumbo a Díaz Ordaz, 150 m, 15 May 1985, F. Ve nt u r a A . 2 14 9 6 (MO, NY).
Tacotalpa, Disperso a la orilla de la selva Milpa em el cerro cerca del Ejido Lázaro Cárdenas al este del ejido, 50 m,
7 June 1979, A. Orozco et al. 2208 (MO). Ve ra cruz - Ll ave: Hidalgotitlán, Hermanos Cedillo, La escuadra por el
Rio Soloxuchil, 150 m, 17 August 1974, M. Vasquez 927 (M). Minatitlán, 2.5 km al N de Poblado 10 por el camino
a Pob. 9, 180 m, 9 May 1986, T. Wendt et al. 5280 (MO).
Taxon omi c not es: Tynanthus guatemalensis is morphologically similar to the Atlantic Forest species T.
micranthus, with which it shares elliptic leaflets with caudate-mucronate apices, lax and conical inflorescences, a
similar corolla length (0.5–0.9 mm in both species), and winged fruits. However, T. guatemalensis presents simple
tendrils (versus trifid in T. micranthus), foliaceous prophylls of the axillary buds (versus minute in T. micranthus)
and a pubescent inflorescence axis (versus glabrescent in T. micrant hus ). Despite the morphological similarities
between those two species, T. guat ema lens is is more closely related to T. cro ati anus, another Central American
species but morphologically very distinct from T. guatemalensis (see notes under T. croatianus).
Nomenclatural notes:—In the protologue of T. guatemalen sis, Donnell Smith (1893) did not clearly indicate
which of the materials examined was the holotype. Two duplicates of the J. Donnell Smith 1488 collection were
located at US, where Donnell Smith worked (Stafleu & Cowan 1985: 698), and the best quality material is here
selected as the lectotype.
7. Tynanthus labiatus (Cham.) Miers (1863: 193). Bignonia labiata Chamisso (1832: 701). Schizopsis labiata
(Cham.) Bureau (1865: 373). Type:—BRAZIL. "Brasil aequinoct.," s.d., F. Sellow s.n. (holotype LE!; isotypes B
destroyed, G-DC! (barcode 133274), HAL! (barcode 98689) photo, K! (barcode 449547), K! (barcode 449547) as
photocopy at MO (2904326), NY! (barcode 579087), US! (barcode 125831)).
Fig. 11: A–E
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FIGURE 11. A–E. Tyna nthu s labi atus : A. Flowering branch; B. Petiolar region showing minute prophylls of the axillary buds; C.
Detail of inflorescence axis showing the indumentum and bracts; D. Flower; E. Fruit (J.R. Pirani 3900, SPF). F–K. T. macr ant hus : F.
Flowering branch; G. Detail of the densely pubescent indumentum in branchlet; H. Interpetiolar glands (R. Lent 42, NY isotype); I.
Flower; J. Open corolla showing the androecium; K. Open calyx showing the gynoecium (A. Jiménez M. 2044, NY). Illustration by
Klei Sousa.
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TAXONOMIC REVISION OF TYNANTHUS (BIGNONIEAE)
Lianas. Branchlets tetragonal to terete, with ritidome (conspicuous when old), finely striated, lenticeled to densely
lenticeled, glabrescent (sometimes pubescent at the nodes), with peltate and patelliform trichomes (sometimes also
with simple); interpetiolar ridge absent or present; interpetiolar patelliform glands absent; prophylls of the axillary
buds 0.8–2.2 mm long, 0.8–1.9 mm wide, minute, shallowly triangular to triangular, pubescent or puberulent to
glabrescent throughout, with simple, peltate and patelliform trichomes. Leaves 2–3 foliolated (more commonly 3);
terminal leaflets often modified into trifid tendrils, without adhesive-disks on tip; petioles and petiolules
glabrescent (sometimes pubescent) throughout surface, with peltate and patelliform trichomes (sometimes also
simple); petioles (0.7–)1.3–6.5 cm long; petiolules 0.6–3.8 cm long; leaflets (3.1–)5.3–13.5 cm long, (1.3–)2.5–7.4
cm wide, membranous to chartaceous, discolor, elliptic; apex acuminate, mucronate; base cuneate or obtuse,
symmetrical; margin entire; the abaxial surface glabrescent (sometimes pubescent) on and near the veins, with
peltate and patelliform trichomes (sometimes also simple); the adaxial surface glabrescent (sometimes pubescent)
on and near the veins, with peltate and patelliform trichomes (sometimes also simple); glandular trichomes
distributed especially on the abaxial surface; second venation weak brochidodromous; pocket domatia with or
without trichomes. Inflorescence axilar or terminal, a thyrse or a compound thyrse, lax, with conical aspect, first
order 5.5–14 cm long, second order 1.5–4 cm long; axis pubescent (at some axis nodes), with simple, peltate and
patelliform trichomes; bracts of the inflorescence predominantly caducous, tomentose to pubescent only at
margins, 0.5–2 mm long; floral bracts 0.4–1.1 mm long; floral pedicels 1–3 mm long. Calyx green to yellowish,
1.6–2.4 mm long, 1.7–2.5 mm wide, with transversal aperture, minutely 5-denticulate (sometimes truncate),
pubescent at teeth to glabrescent outside, with patelliform glands; lobes 0.1–0.3 mm long. Corolla white, cream or
pale yellow (sometimes pale pink), 1–1.4 cm long, 3.5–6 mm wide at the tube opening; tube 4–7 mm long,
internally tomentose at the base, with simple and long and short stipitate trichomes; nectar guides present, yellow;
lobes densely pubescent to pubescent throughout lower ones and at the margin of upper ones; upper ones 0.4–1.4
mm long, 0.9–2.4 mm wide, acute to obtuse; lower ones 1.5–4 mm long, 2.4–4 mm wide, obtuse to rounded.
Androecium with fertile stamens inserted 1.1–2.5 mm from the base of the corolla; shorter ones 2.5–7 mm long;
longer ones 3.5–9 mm long; anthers thecae 1–1.4 mm long, obovate to elliptic, subexserted; connective extending
0.2–0.3 mm beyond anther attachment; staminode 1.1–2.5 mm long, glabrescent, with long and short stipitate
trichomes. Gynoecium 4–12 mm long; ovary 1–1.2 mm long, 0.7–0.8 mm wide, conical, velutinous; style 3–10 mm
long, tomentose to pubescent at the base. Fruit a linear flattened capsule, 10–27 cm long, 2.4–3.1 cm wide, woody,
smooth to granular near the midvein and granular near the margins, without lenticels, glabrescent, with peltate and
patelliform trichomes; central ridge single, slightly or not prominent; margins prominently raised (winged), 0.6–1.2
cm wide. Seeds body 0.9–1 cm long, 0.7–0.8 cm wide; wings not seen.
Phenology:—Flowers and produces fruits from January to May.
Distribution and habitats:—Occurs mainly in moist broadleaf forests, although some populations also reach
dry forests from Brazil (Bahia, Minas Gerais and Rio de Janeiro) (Fig. 12).
Additional specimens examined:—BRAZIL. Sin loc., s.d., A. Glaziou 16269 (P). Bahia: Igreja Velha, 1841,
J.S. Blanchet 3267 (BM, BR, F, G, G-DC, LE, M, NY, P). Caetité, Café Baiano, 9 km E de Caetité em direção a
Brumado, 800 m, 7 March 1994, V.C. Souza et al. 5334 (SPF 108378, fruit attached to an Adenocalymma Mart. ex
Meisn. specimen). Rui Barbosa, Serra do Orobó, Fazenda Bom Jardim, 426 m, 26 May 2005, L.P. de Queiroz et al.
10661 (SPF). Minas Gerais: Sin loc., s.d., E.P. Heringer s.n. (RB barcode 58263). Catas Altas?, 1882?, A. Glaziou
14108 (BR, F, G, K, LE, MO, P, US). Coronel Pacheco, Fazenda Companhia, 29 February 1944, E.P. Heringer
1331 (RB). Monjolos, Estrada Corinto–Conselheiro Mata, margem esquerda do Rio Pardo Pequeno, 540 m, 19
February 2011, M.C. Medeiros & R.B. Louzada 31 (SPF). Santo Hipólito, Estrada Corinto–Conselheiro Mata, a 6
km de Santo Hipólito, 550 m, 4 April 1996, J.R. Pirani et al. 3775 (SPF); 12 January 1998, J.R. Pirani et al. 3900
(SPF); 19 February 2011, M.C. Medeiros & R.B. Louzada 30 (SPF); Cerca de 5 km além de Santo Hipólito em
direção a Monjolos, no antigo leito da estrada de ferro, afloramento de calcáreo à margem esquerda do Rio Pardo
Pequeno, 500 m, 24 March 1997, R. Mello-Silva et al. 1317 (SPF). Volta Grande, Foz do rio angu, January 2007,
L.C.S. Viana & G.M. Maciel s.n. (BHCB 109098). Rio de Janeiro: Sin loc., 1832, L. Riedel 88 (LE, NY). Vallée du
Rio Comprido, 18 February 1871, A. Glaziou 4703 (P). Itaguaí, Rio Mazomba, 12 January 1950, L.E.M. Filho
1029 (R); 12 January 1950, A.C. Brade & A. Duarte 20165 (MO, NY, RB). Rio de Janeiro, Parque do Jardim
Botânico do Rio de Janeiro, Pedra do Marinheiro, 25 January 1989, R. Marquete et al. 202 (RB); Ibid., trecho entre
a caixa d’água do Caminho do Boi até Pedra do Marinheiro, 5 March 1991, M. Madruz et al. 678 (K).
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FIGURE 12. Distribution of Tyna nth us lab iat us.
Taxonomic notes:Tyn anthus labiatus can be easily recognized in the field by the conspicuous ritidome
encountered in old branches and stem. Apart from that, T. labiatus is characterized by the glabrescent elliptic
leaflets, conical inflorescences and winged fruits, all of which are also shared with the Amazonian T.
schumannianus. However, T. la biatu s has glabrous inflorescences axes, pubescent only at nodes (versus densely
pubescent to pubescent inflorescences axes throughout in T. schumann ianus) and larger corollas, 1–1.4 cm (versus
smaller corollas, 0.4–0.9 cm in T. schumannianus). Tynanthus labiatus is a close relative of T. fasciculatus
(Medeiros & Lohmann 2015; see T. fasci culatus notes).
8. Tynanthus macranthus Williams (1967: 250) (as “Tynnanthus”). Type:—COSTA RICA. Heredia: Roadside 8
km south of San Miguel, 9 July 1964, R.W. Lent 42 (holotype F! (1622130); isotypes NY! (barcode 328981), OKL!
(barcode 01 0097215) photo, US! (barcode 125787)).
Fig. 11: F–K
Lianas. Branchlets terete, without ritidome, finely striated, densely lenticeled, densely puberulent, with simple and
peltate trichomes; interpetiolar ridge absent or present; interpetiolar patelliform glands present; prophylls of the
axillary buds 1–3 mm long, 1–2.5 mm wide, minute, shallowly triangular, densely puberulent throughout, with
simple and peltate trichomes. Leaves 2 foliolated; terminal leaflets often modified into simple tendrils, without
adhesive-disks on tip; petioles and petiolules densely puberulent throughout surface, with simple, peltate and
patelliform trichomes; petioles (0.6–)0.8–2.3 cm long; petiolules (0.3–)0.7–2 cm long; leaflets (2.6–)4.5–14.9 cm
long, (1–)2.7–6.4 cm wide, chartaceous to coriaceous, discolor, elliptic; apex acuminate, mucronate; base cuneate
or obtuse, symmetrical; margin entire; the abaxial surface densely puberulent to puberulent throughout (sometimes
only on and near the veins), with simple, peltate and patelliform trichomes; the adaxial surface puberulent to
glabrescent on and near the veins, with simple and peltate trichomes; glandular trichomes distributed especially on
the abaxial surface; second venation weak brochidodromous; pocket domatia with or without trichomes.
Inflorescence axilar or terminal, a thyrse, dense, with corymbose or subcorymbose aspect, 2.5–3–6 cm long; axis
densely puberulent, with simple and peltate trichomes; bracts of the inflorescence predominantly caducous,
densely puberulent throughout, 1–2 mm long; floral bracts 0.8–1.2 mm long; floral pedicels 2–6 mm long. Calyx
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TAXONOMIC REVISION OF TYNANTHUS (BIGNONIEAE)
green, 5–9(–10) mm long, 4–5 mm wide, with transversal aperture, minutely 5-denticulate (sometimes truncate),
densely puberulent throughout outside, without patelliform glands; lobes 0.2–0.5 mm long. Corolla white, 2–3.8
cm long, 2.5–4 mm wide at the tube opening; tube 5–11 mm long, internally tomentose at the base, with simple and
long and short stipitate trichomes; nectar guides present, yellow; lobes densely pubescent to pubescent throughout
lower ones and at the margin of (sometimes throughout) upper ones; upper ones 2–8 mm long, 1.5–4 mm wide,
obtuse to rounded (sometimes acute); lower ones 4–8(–10) mm long, 3–7 mm wide, obtuse to rounded.
Androecium with fertile stamens inserted ca. 3 mm from the base of the corolla; shorter ones 7–13 mm long; longer
ones 9–18 mm long; anthers thecae 1.7–2.5 mm long, elliptic, inserted; connective extending 0.2–0.4 mm beyond
anther attachment; staminode 4–7 mm long, glabrescent, with long and short stipitate trichomes. Gynoecium 14–22
mm long; ovary 1.8–2.5 mm long, 0.9–1.1 mm wide, conical to oblong, densely pubescent; style 12–19 mm long,
densely pubescent throughout. Fruit not seen. Seeds not seen.
Phenology:—Flowers from April to November; the fruiting season is unknown.
Distribution and habitat:—Occurs in moist broadleaf forests from Costa Rica (Heredia and Limón) and
Panama (Veraguas) (Fig. 13).
FIGURE 13. Distribution of Tyna nth us mac ran thu s.
Additional specimens examined:—COSTA RICA. Heredia: Entre San Miguel y Cariblanco de Sarapiqui,
725 m, 10 July 1964, A. Jiménez M. 2044 (F photo, NY). Limón: Talamanca, Between Cahuita and the oil drilling
platforms beyond Suretka, 100–300 m, 23–25 April 1982, K. Barringer et al. 2671 (F, MO). PANAMA. Veraguas:
Vicinity of Continental Divide, third branch of Rio Santa Maria to drop-off to lowlands, 12–15 km NW of Santa
Fe, 650–750 m, 16–17 November 1974, R. Dressler 4846 (MO). NW of Santa Fe, 8.8 km from Escuela Agricola
Alto de Piedra, 17 May 1975, R. Dressler et al. 6191 (MO, US).
Taxonomic notes:Tynanthus macranthus is easily distinguished from the other species of the genus by the
large flowers (2–3.8 cm), with conspicuously bilabiate corolla due to the short corolla tube (0.5–1.1 cm). Apart
from that, T. macranthus is also characterized by the densely puberulent branchlets, simple tendrils, interpetiolar
patelliform glands and corymbose to subcorymbose inflorescences. Tynanthus macranthus was the first species to
diverge in Tynanthus (Medeiros & Lohmann 2015) and is sister to the remaining species of the genus.
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9. Tynanthus micranthus Mello ex Schumann (1894: 221) (as "Tynnanthus micrantha"). Lectotype (designated by
Lohmann, in Lohmann & Taylor 2014: 469):—BRAZIL. São Paulo: Campinas, 30 September 1867, J.C. de Mello 26
(P! (barcode 481491); isolectotypes F! (999019), K! (barcode 449546), NY!, P! (barcodes 481488, 481489, 481490),
P! (barcode 481489) as photocopy at MO, UPS! as photocopy at K, US! (2515396)).
Fig. 14: A–D
FIGURE 14. A–D. Tyn ant hu s mi cr an th us : A. Flowering branch; B. Open corolla showing the androecium; C. Open calyx showing
the gynoecium (M.C. Medeiros 32, SPF); D. Fruit (J.S. Carneiro 300, FUEL). E–I. T. p an ure ns is : E. Branch; F. Interpetiolar region
showing the foliaceous prophylls of the axillary buds; G. Inflorescence; H. Flower; I. Fruit (E–F and I: A.H. Gentry 21066, MO)(G–H:
G. Klug 1972, NY). Illustration by Klei Sousa.
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TAXONOMIC REVISION OF TYNANTHUS (BIGNONIEAE)
Lianas. Branchlets tetragonal to terete, with or without ritidome, finely striated, lenticeled to densely lenticeled,
glabrescent, with peltate trichomes; interpetiolar ridge absent (sometimes present); interpetiolar patelliform glands
absent; prophylls of the axillary buds 0.8–1.6 mm long, 0.5–1.2 mm wide, minute, triangular to shallowly
triangular, glabrous. Leaves 2–3 foliolated (more commonly 3); terminal leaflets often modified into trifid tendrils,
without adhesive-disks on tip; petioles and petiolules glabrescent throughout surface (sometimes pubescent at the
junction of petiolules), with peltate trichomes (sometimes also with simple); petioles (0.5–)1.5–5.3 cm long;
petiolules (0.3–)0.6–2.7 cm long; leaflets (2.3–)4.1–9.1 cm long, (0.7–)1.3–4.2 cm wide, membranous to
chartaceous, discolor or concolor, elliptic; apex caudate, mucronate; base cuneate or obtuse, symmetrical; margin
entire; the abaxial surface glabrescent, with peltate and patelliform trichomes; the adaxial surface glabrescent, with
peltate trichomes; glandular trichomes distributed especially on the abaxial surface; second venation weak
brochidodromous; pocket domatia with (sometimes without) trichomes. Inflorescence axilar or terminal, a thyrse or
a compound thyrse, lax, with conical aspect, first order 3.5–5–13.5 cm long, second order 1.5–3 cm long; axis
glabrescent, with peltate trichomes; bracts of the inflorescence predominantly caducous, glabrescent (sometimes
pubescent only at margins), 0.7–6(–9) mm long; floral bracts 0.5–1 mm long; floral pedicels 1–4 mm long. Calyx
green, 1.2–2 mm long, 1.5–2.3 mm wide, with transversal aperture, minutely 5-denticulate (sometimes truncate),
glabrescent (sometimes pubescent at teeth) outside, with patelliform glands; lobes 0.1–0.6 mm long. Corolla white,
cream or pale yellow (sometimes pale green), 0.5–0.9 cm long, 2–3.5 mm wide at the tube opening; tube 2.5–3.5
mm long, internally tomentose to pubescent at the base or glabrescent, with simple and long and short stipitate
trichomes; nectar guides absent, but with a path of long and short stipitate trichomes; lobes densely pubescent to
pubescent throughout lower ones and at the margin of upper ones; upper ones 0.4–1.3 mm long, 0.7–1.4 mm wide,
acute to obtuse; lower ones 2–3.1 mm long, 1.5–2.7 mm wide, obtuse to rounded. Androecium with fertile stamens
inserted 0.8–1 mm from the base of the corolla; shorter ones 2.1–3.1 mm long; longer ones 2.6–4.5 mm long;
anthers thecae 0.6–0.8 mm long, obovate to elliptic, subexserted; connective extending ca. 0.2 mm beyond anther
attachment; staminode 1–1.2 mm long, with long and short stipitate trichomes. Gynoecium 4.1–4.6 mm long; ovary
0.9–1 mm long, 0.5–0.6 mm wide, conical, velutinous; style 3–3.7 mm long, tomentose at the base. Fruit a linear
flattened capsule, 7.2–16.5 cm long, 0.9–1.3 cm wide, coriaceous to woody, smooth to granular near the midvein
and granular near the margins, without lenticels to densely lenticeled, glabrescent (sometimes pubescent at
extremities), with peltate and patelliform trichomes (sometimes also with simple); central ridge single, not
prominent; margins prominently raised (winged), 0.1–0.4 cm wide. Seeds body 1.2–1.4 cm long, 0.6–0.8 cm wide;
wings 0.4–1 cm long.
Phenology:—Flowers from September to December and produces fruits from December to May.
Distribution and habitat:—Occurs in moist broadleaf forests from Brazil (Mato Grosso do Sul, Paraná and
São Paulo) and Paraguay (Alto Paraná and Canindeyú) (Fig. 15).
Additional specimens examined:—BRAZIL. Reserva Florestal II, 21 October 2005, Adriana et al. s.n.
(HRCB 42809). Mato Grosso do Sul: Iguatemi, Arredores, 400 m, 21 October 1987, G. H a t s c h b a c h & J . M . S i l v a
51518 (BR, MO, US). Paraná: Parque Nacional do Iguú, N border, 2 1/2 km W of road to Capanema, S of
Jardinópolis, 23 November 1966, J.C. Lindeman & J.H. de Haas 3354 (MO, US); Ibid., by forest road near
Aranha, SE of Jardinópolis, 24 November 1966, J.C. Lindeman & J.H. de Haas 3409 (F photo, K, MO, NY, P, RB,
US). Alvorada do Sul, Fazenda Ingá, 19 May 2000, O.C. Pavão s.n. (FUEL 29345). Arapongas, Fazenda Solana,
26 April 2004, J.S. Carneiro et al. 300 (FUEL, HRCB). Califórnia, Sítio Ogido, 9 November 1986, S.N. Ogido 7
(FUEL); Rio Água do Oito, 19 October 2000, O.C. Pavão 5 (FUEL). Cambé, Mata da Morada do Sol, 8 October
1999, C.A. Avanzi et al. s.n. (FUEL 29323). Campo Mourão, Rio da Várzea, 515 m, 14 October 1965, G.
Hatschbach 13033 (F, K, NY, P, UPCB, US). Céu Azul, Boa Vista, 22 October 1969, G. Hatschbach 22593 (F, K,
MO, NY, UPCB); Parque Nacional do Iguaçú, trilha da Jacutinga, 658 m, 12 October 2011, J.A. Lombardi et al.
8739 (HRCB). Congonhinhas, Fazenda Planalto, beira da represa, 21 October 1998, A.L. Cavalheiro et al. s.n.
(FUEL 23840, RB barcode 535433, SPF barcode 197643, SPSF 40442). Dois Vizinhos, Região compreendida
entre os municípios de Dois Vizinhos e Pato Branco, proximidades do rio Chopim, 500 m, 8 December 2001, P.H.
Labiak & M. Kaehler 2016 (BHCB). Douradina, Fazenda Santa Rosa?, 2 November 1959, R. Braga & R. Lange 80
(UPCB, US). Fênix, Fazenda Água Azul, 1 November 1998, J.M. Silva et al. 2587 (ESA, FUEL, G, INPA, K, R,
RB, SP, SPF, UPCB). Goioerê, Parque Municipal Antônio Sestak, 25 September 2007, E.M. Silva s.n. (RB barcode
527993). Ibiporã, Sítio São José, Água das Abóboras, 19 October 1997, E.M. Nakano & I.M. Medri s.n. (FUEL
24556). Londrina, Floresta dos Irmãos Godoy, 2 October 1985, F. C . S i l v a e t a l . 9 0 1 (FUEL, MO); Ibid., borda do
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extremo norte do Parque, próximo ao rio, 600 m, 12 October 2011, M.C. Medeiros & E.F. Rossetto 32 (SPF);
Fazenda Santa Ana, 24 October 1985, A.O.S. Vieira et al. s.n. (FUEL 1661); Ibid., 24 October 1985, A.O.S. Vieira
et al. 41 (MO); Rio 3 Bocas, Sítio do Gasparino, 22 October 1986, C. Zampieri et al. 41 (FUEL, HRCB, R);
Proximidades do EMAUS, 27 September 1997, M.C.B. Azevedo et al. s.n. (FUEL 22931); Estância Cabral, 10
October 1999, S.F. Andrade et al. s.n. (HRCB 55991); Fazenda Figueira-Paiquerê, Fragmento 1, December 2002,
M.C. Lovato et al. 381 (FUEL); Fazenda Escola, 3 March 2009, G. F. S a n t o s 7 (FUEL); Parque Municipal Arthur
Thomas, trilha da Capivara, 520 m, 14 October 2011, M.C. Medeiros & E.F. Rossetto 34 (SPF). Lupionópolis, Mata
São Pedro, 6 May 2005, J.S. Carneiro et al. 382 (FUEL). Matelândia, 13 October 1962, G. Hatschbach 9342 (K).
Nova Prata do Iguaçu, Rodovia Nova Prata a Cruzeiro do Iguaçu, Rio Jacaratiá, 15 October 1997, J.M. Silva et al.
2214 (ESA, G, HRCB, SPF). Rolândia, Sítio Jevy, Contorno Norte, 15 October 1996, R. Losi et al. s.n. (FUEL
25345). São Pedro do Ivaí, Fazenda Barbacena, 16 October 2003, O.S. Ribas et al. 5527 (ESA, FUEL, G, MO, RB).
Telêmaco Borba, margem da estrada de acesso ao eixo da barragem, 680 m, 22 September 2008, M. Kaehler 277
(UPCB); Desmatamento para construção do Eixo da Barragem da UH Mauá, 680 m, 23 September 2008, M.
Kaehler 310 (UPCB). São Paulo: Adamantina, Sítio São Paulo estrada 6, 7 September 1995, E.Y. Sasaki s.n. (ESA
103223, FUEL 29302, SPF barcode 201517, VIC 31746). Gália, Estação Ecológica dos Caetetus, 12 February
2003, R. Udulutsch & R. Tsuji 1619 (HRCB). Jundiaí, Reserva Biológica Municipal da Serra do Japi,
Base–Mirante–estrada para DAE–Base, 850–985 m, 23 October 2007, J.A. Lombardi & S.M. Hieda 6990 (HRCB).
Monte Alegre do Sul, Fazenda Benati, 17 March 1995, L.C. Bernacci et al. 1366 (HRCB, SPF). Piracicaba, Mata
da Pedreira, ESALQ, Beira do Ribeirão Piracicamirim, 2 October 1985, E.L.M. Catharino 444 (ESA, SP). Rio
Claro, Fazenda São José, 30 October 2001, R. Udulutsch et al. 423 (HRCB); Ibid., 7 October 2002, M.A. Assis &
A.G. Manzatto 1599 (HRCB). PARAGUAY. Alto Paraná: Reserva Biológica Limoy, cerca del río Limoy y
Embalse de la Represa Itaipú em el río Paraná, 14 October 1996, A. Schinini et al. 31364 (G, MO). Canindeyú:
Circa Guadelupe, Canendiyu, 27 October 1978, L. Bernardi 18227 (BM, F, G, MO, NY).
FIGURE 15. Distribution of Tyna nth us mic ran thu s.
Taxonom ic n ote s:Tynanthus micranthus is characterized by the glabrescent branchlets and leaflets, lax
inflorescences, small corollas (0.5–0.9 cm), and winged fruits. This species is morphologically similar to the
Central American T. guatemal ensi s, from which it differs on some morphological features (see taxonomic notes
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TAXONOMIC REVISION OF TYNANTHUS (BIGNONIEAE)
under T. guatemalensis). Tyn anthus micra nthus is sister to a clade that includes two other Atlantic Forest species, T.
fasciculatus and T. labiatus (Medeiros & Lohmann 2015). Morphologically, T. micranthus is most similar to T.
labiatus, with which it shares glabrescent branchlets and leaflets; however, these two species can be separated by
the smaller flowers of T. micranthus (0.5–0.9 cm versus 1–1.4 cm in T. labiatus).
10. Tynanthus panurensis (Bureau) Sandwith (1953: 465) (as “Tynnanthus”). Schizopsis panurensis Bureau (1865:
373). Lectotype (designated here):—BRAZIL. Amazonas: “Prope Panuré ad Rio Uaupés”, October 1852–January
1853, R. Spruce 2626 (P! (barcode 468600); isolectotypes BM! (barcode 992358), BM! (barcode 992358) as
photocopy at MO (2300302), BR! (barcode 876218), E! (barcode 394578) photo, G!, G! as photocopy at F (785147),
G! as photocopy at MO (1693115), G! as photocopy at NY, G! as photocopy at US, K! (barcode 449548), LE!, P!
(barcodes 3606843, 608100), TCD! (barcode 550) photo).
Fig. 14: E–I
Lianas. Branchlets tetragonal to terete, without ritidome, finely striated, lenticeled to densely lenticeled, puberulent
to glabrescent, with simple and peltate trichomes; interpetiolar ridge absent or present; interpetiolar patelliform
glands absent; prophylls of the axillary buds 2.5–5 mm long, 1.5–3.5 mm wide, foliaceous, obovate, puberulent
throughout, with simple, peltate and patelliform trichomes. Leaves 2–3 foliolated (more commonly 3); terminal
leaflets often modified into trifid tendrils, without adhesive-disks on tip; petioles and petiolules puberulent to
glabrescent throughout surface, with simple and peltate trichomes; petioles 3–12 cm long; petiolules 0.7–7.8 cm
long; leaflets (3.7–)6.5–20.4 cm long, (2–)4.1–13.7 cm wide, membranous to chartaceous or coriaceous, discolor,
elliptic to ovate; apex acuminate, mucronate; base cuneate or obtuse, symmetrical or asymmetrical; margin entire;
the abaxial surface pubescent to glabrescent on and near the veins, with simple, peltate and patelliform trichomes;
the adaxial surface pubescent to glabrescent on and near the veins, with simple and peltate trichomes; glandular
trichomes distributed especially on the abaxial surface; second venation weak brochidodromous; pocket domatia
without trichomes. Inflorescence axilar or terminal, a thyrse, lax, with conical aspect, (3–)5–13 cm long; axis
pubescent to puberulent, with simple and peltate trichomes; bracts of the inflorescence caducous or persistent,
pubescent throughout, 1–2.1 mm long; floral bracts 0.6–0.9 mm long; floral pedicels (0.5–)1–9 mm long. Calyx
green, 2.5–4 mm long, 1.9–3 mm wide, with transversal aperture, truncate or minutely 5-denticulate, puberulent
throughout outside, without patelliform glands; lobes 0.1–0.3(–0.6) mm long. Corolla cream or pale yellow
(sometimes pale lilac), 1.2–1.7 cm long, 4–6 mm wide at the tube opening; tube 6–7 mm long, internally tomentose
at the base, with simple and long and short stipitate trichomes; nectar guides absent, but with a path of long and
short stipitate trichomes; lobes densely pubescent to pubescent throughout lower ones and at the margin of upper
ones; upper ones 0.6–1.4 mm long, 0.9–1.9 mm wide, acute to obtuse; lower ones 3.2–5.1 mm long, 3.1–4.2 mm
wide, obtuse to rounded. Androecium with fertile stamens inserted 1.5–3 mm from the base of the corolla; shorter
ones 3.5–6 mm long; longer ones 4–8 mm long; anthers thecae 1–1.2 mm long, obovate to elliptic, subexserted;
connective extending 0.2–0.4 mm beyond anther attachment; staminode 3–3.3 mm long, glabrous. Gynoecium
7–13 mm long; ovary 1.5–2 mm long, 0.7–0.9 mm wide, conical, densely pubescent; style 5–12 mm long,
tomentose to pubescent at the base. Fruit a linear flattened capsule, 22–25 cm long, (0.5–)0.9–1.4 cm wide,
coriaceous to woody, smooth to granular near the midvein and granular near the margins, densely lenticeled,
pubescent, with simple, peltate and patelliform trichomes; central ridge single, not prominent; margins slightly
raised (unwinged), 0.1–0.2 cm wide. Seeds not seen.
Phenology:—Flowers from October to April. Two fruiting collections were observed, one made in December
and the other in May (fallen fruits).
Distribution and habitat:Occurs in moist broadleaf forests from Brazil (Amazonas), Colombia
(Amazonas and Caquetá), Ecuador (Orellana) and Peru (Loreto, Pasco and San Martín) (Fig. 16).
Additional specimens examined:—BRAZIL. Amazonas: SEPLAC, km 60 on Manaus–Caracarai Road (BR
174), 1 December 1974, A.H. Gentry 12997 (INPA photo, MG, MO, R). COLOMBIA. Amazonas: Araracuara,
Villa Azul, Rio Caquetá, margen izquierda frente isla Sumaeta, 12 May 1989, C. Londoño et al. 625 (MO); Ibid.,
200–300 m, 22 November 1989, C. Londoño et al. 1449 (MO). Caquetá: 10 km S of San Jose de Fragua, 320 m,
11 January 1974, A.H. Gentry et al. 9144 (MO). Araracuara, 28 January 1989, A.H. Gentry et al. 65294 (MO).
ECUADOR. Orellana: Tiputini Biodiversity Station, Río Tiputini, about 20 km (via air) east of confluence with
Río Tivacuno, Near Erwin Plot #1 off Sendero Harpia, 250 m, October 1998, R.J. Burnham et al. 2307 (MO).
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PERU. Loreto: Alto Amazonas, Capihuari, 5 km NE of Andoas on Río Capihuari, near Ecuador border, Along oil
pipeline, 240 m, 17 November 1979, A.H. Gentry & C. Díaz 28174 (F, MO, NY); Few km W of Shucushayacu on
Rio Huallaga above Yurimaguas, 180 m, 11 October 1985, A.H. Gentry et al. 52229 (SPF). Mariscal Ramón
Castilla, Pebas, Bora Native community of Brillo Nuevo, Yaguasyacu River (affluent of Ampiyacu River), ca. 150
km ENE of Iquitos, 106 m, 31 October 1981, J. Treacy & J.B. Alcorn 259 (F); 4 November 1981, J. Treacy & J.B.
Alcorn 333 (F). Maynas, Florida, Rio Putumayo, at mouth of Rio Zubineta, 200 m, March–April 1931, G. K l u g
1972 (BM, F, MO, NY, US); Nanay to (ad) Iquitos, 200 m, 10 December 1958, F. Woytkowski 5151 (K, MO); Río
Ampiyacu, Pebas and vicinity, Trail north from town, 10 April 1977, T. Plowman et al. 6706 (F photo, MO); Near
Brilla Nueva, Borro Indian village on upper Rio Yaguasyacu, tributary of Rio Ampiyacu, 8 November 1977, A.H.
Gentry & J. Revilla 20476 (F); Mishana, Rio Nanay, 130 m, 3 December 1977, A.H. Gentry et al. 21066 (F, MO,
NY); Ibid., Río Nanay halfway between Iquitos and Santa Maria de Nanay, Cerca del campamento No. 1, 140 m,
19 May 1979, C. Diaz et al. 1105 (F, MO); Ecuador border, 1–2 km from Puerto Peru (military post 8 km from
mouth of Río Gueppi, tributary of Río Putumayo), 200 m, 18 May 1978, A.H. Gentry et al. 22055 (MO);
Pucaurquillo, Trail behind Bora villiage towards mounte, 18 August 1981, R. Hahn & R. Tredwell 3I? (MO
2927651); Rio Ampiyacu, Pucuarquillo, Path to Witoto central forest, behind Pucuarquillo, 21 September 1981, R.
Hahn et al. 145 (MO); Iquitos, Allpahuayo, Estación Experimental del Instituto de Investigaciones de la Amazonia
Peruana (IIAP), Muestreo de 1000 m2, Transecto No 9, 150–180 m, 26 May 1991, R. Vásquez & N. Jaramillo
16543 (MO); Ibid., Linea F de la parcela de 25 ha, subparcela 5, 150 m, 20 March 1992, R. Vásquez et al. 17806
(MO); San Francisco de Orellana, Rio Napo, trail of Pue San Pedro below caserio of Juancho Playa, 9 November
1978, M. Rimachi Y. 4064 (F photo, MO, NY). Requena, Jenaro Herrera, Río Ucayali below Requena, 9 December
1977, A.H. Gentry et al. 21323 (F, MO); Ibid., Río Ucayali, 19 February 1987, A.H. Gentry et al. 56126 (F, MO);
Sapuena, Basin of Río Ucayali, Arboretum Jenaro Herrera, 21 April 1987, D.C. Daly et al. 5122 (MO, NY). Pasco:
Oxapampa, Cabeza de Mono, Río Iscozacin, 10 km SW of Iscozasin, Palcazu Valley, 320 m, 9 June 1983, A.H.
Gentry et al. 41745 (MO); Shiringamazu, ca 20 km S of Iscozacin, Río Palcazu Valley, 300 m, 6 July 1988, A.H.
Gentry et al. 63339A (MO). San Martín: San Martin, Near km 50 Yurimaguas–Tarapoto road, 200 m, 12 October
1985, A.H. Gentry et al. 52280 (F, MO).
FIGURE 16. Distribution of Tyna nth us pan urens is.
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TAXONOMIC REVISION OF TYNANTHUS (BIGNONIEAE)
Taxo no mi c no te s: Tynanthu s pan urensis is characterized by the foliaceous prophylls of the axillary buds,
unusually long petioles (up to ca. 12 cm) and petiolules (up to ca. 7.8 cm), lax inflorescences, corollas 1.2–1.7 cm
long and fruits with slightly raised margins. This species is closely related to T. pubescens (Medeiros & Lohmann
2015), with which it shares lax inflorescences and a similar corolla length (1.2–1.7 cm in T. panurensis and 1–1.6
cm in T. pubescens); however, T. pan urensis can be separated by the foliaceous prophylls of the axillary buds
(versus minute prophylls in T. pubesce ns), and only slightly raised fruit margins (versus prominently raised in T.
pubescens). Despite the phylogenetic proximity between these species, T. panurensis is morphologically most
similar to T. densif l o rus, with which it shares some morphological features (see taxonomic notes under T.
densiflorus).
Nomenclatural notes:—Three duplicates of the collection Spruce 2626 were located at P, and the best quality
flowering material is here selected as the lectotype.
11. Tynanthus polyanthus (Bureau) Sandwith (1953: 465) (as “Tynnanthus”). Schizopsis polyantha Bureau (1865:
378). Lectotype (designated by Lohmann, in Lohmann & Taylor 2014: 469):—PERU. San Martín: “Prope Tarapoto”,
1855–1856, R. Spruce 4895 (P! (barcode 481492); isolectotypes BM! (barcode 603754), BR! (barcode 876198), C!
photo, C! as photocopy at F (718222), C! as photocopy at MO (1693114), E! (barcode 259199) photo, F! (1012196)
(fragments), F! (875569) photo, G!, G! as photocopy at F (785146), G! as photocopy at MO (1693113), G! as
photocopy at NY, G! as photocopy at US, K! (barcodes 202056, 202057, 202058, 202059), LE!, MPU! (barcode
16048) photo, NY! (barcode 328924), P! (barcode 481493, 481494), S! (04-3493) photo, TCD! (barcode 549) photo).
Schizopsis polyantha var. boliviana Bureau (1865: 379). Type:BOLIVIA. La Paz:Environs de Mururata, 1839, J.B.
Pentland 33 (holoty pe P! (barcode 608099)).
Tyn an thus m yriant hu s Bureau & Schumann (1896: 197) (as “Tyn n a nthus). Lectotype (designated here):—PERU. Loreto: “Ad
Yu ri m ag u as ” , A pr i l 18 3 1, E.F. Poeppig 2388 (W, W! as photocopy at F (956391), W! as photocopy at K, W! as photocopy
at MO (1693116), W! as photocopy at US; isolectotypes F! (875526), LE!).
Tynanthus lindmanii Schumann (in Bureau & Schumann 1896: 409) (as Tynna n t h u s ). Lectotype (designated
here):—BRAZIL. Mato Grosso: “Barra do Rio dos Bugres” (Barra do Bugres), 26 February 1894, C.A.M. Lindman A2967
(S! (09-21559) photo; isolectotypes UPS! photo, S! (09-21560) photo, S! (09-21560) as photocopy at K).
Tyn an thus ca ryophy ll eu s (Bello) Alain (1965: 352) (asTynna n t h us). Bignonia? caryophyllea Bello (1881: 293). Neotype
(designated here):—PORTO RICO: “Forest near Humacao”, 28 February 1926, N.L. Britton 8614 (NY! (barcode 874601);
isoneotype US! (barcode 477379)).
Tynanthus villosus Gentry (1976: 60) (as “Tynna nthu s). Type:—PERU. San Martín: Tocache Nuevo, Quebrada de Canuto, 18
August 1973, J. Schunke Vigo 6852 (holotype MO! (2175155); isotypes F! (1763392), K! (barcode 449538), RB!
(barcodes 536855, 536925)). syn nov.
Fig. 17: A–F
Lianas. Branchlets conspicuously tetragonal (with salient extremities) to terete, with or without ritidome, finely
striated (sometimes not evident, when young), few to densely lenticeled (sometimes without lenticels), villous to
puberulent or glabrescent, with simple and peltate trichomes; interpetiolar ridge absent or present; interpetiolar
patelliform glands absent; prophylls of the axillary buds 4–19(–25) mm long, 3–18(–23) mm wide, foliaceous,
ovate or obovate to elliptic, villous to puberulent throughout, with simple, peltate and patelliform trichomes.
Leaves 2–3 foliolated (more commonly 2); terminal leaflets often modified into simple tendrils, without adhesive-
disks on tip; petioles and petiolules villous to puberulent throughout surface, with simple and peltate trichomes;
petioles (0.9–)1.4–5 cm long; petiolules (0.2–)0.6–4.6 cm long; leaflets (2.5–)4–16.5 cm long, (1.6–)2.5–12.6 cm
wide, membranous to chartaceous (sometimes subcoriaceous), discolor or concolor, elliptic to ovate; apex
acuminate, mucronate; base cuneate, obtuse, truncate or subcordate, symmetrical or asymmetrical; margin entire
(rarely dentate); the abaxial surface villous to puberulent (sometimes glabrescent) throughout (sometimes only on
and near the veins), with simple, peltate and patelliform trichomes; the adaxial surface villous to pubescent
(sometimes glabrescent) throughout (sometimes only on and near the veins), with simple, peltate and patelliform
trichomes; glandular trichomes distributed especially on the adaxial surface (sometimes evenly distributed
throughout both surfaces); second venation weak brochidodromous; pocket domatia with or without trichomes.
Inflorescence axilar or terminal, a thyrse or a compound thyrse, lax, with conical aspect, first order 3.5–15 cm long,
second order 1.5–8 cm long; axis villous to pubescent, with simple and peltate trichomes; bracts of the
inflorescence caducous or persistent, villous to pubescent throughout, 0.5–3.3(–6) mm long; floral bracts 0.4–0.7
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Phytotaxa 216 (1) © 2015 Magnolia Press
FIGURE 17. A–F. Tynanthus polyanthus: A. Flowering branch; B. Interpetiolar region showing the foliaceous prophylls of the
axillary buds; C. Detail of inflorescence axis showing bracts; D. Detail of petiolule showing the upper canalicule; E. Flower (M.C.
Medeiros 40, SPF); F. Fruit showing the flattened cross section (P. Núñez 11199, MO). G–K. T. pu besc ens: G. Flowering branch; H.
Detail of pubescent indumentum in branchlet (S. Mori 23971, NY); I–J. Interpetiolar glands; K. Fruit (A.H. Gentry 25579, F).
Illustration by Klei Sousa.
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TAXONOMIC REVISION OF TYNANTHUS (BIGNONIEAE)
mm long; floral pedicels 0.5–7.3(–10.5) mm long. Calyx green to yellowish, 1–2.7 mm long, 1.2–2.2 mm wide,
with transversal (sometimes oblique) aperture, minutely 5-denticulate (sometimes truncate), tomentose to
puberulent throughout outside, without patelliform glands; lobes 0.1–0.3(–0.6) mm long. Corolla white, cream or
pale yellow (sometimes pale lilac, pale red or pale blue), 0.4–0.8 cm long, 1.2–3.5 mm wide at the tube opening;
tube 2–4.5 mm long, internally glabrescent, with long and short stipitate trichomes; nectar guides absent, but with a
path of long and short stipitate trichomes; lobes densely pubescent to pubescent throughout (sometimes only at
margins of) lower ones and at margins of or throughout upper ones; upper ones 0.4–1.6 mm long, 0.5–1.5 mm
wide, acute to obtuse; lower ones 1.3–3 mm long, 1.2–2.8 mm wide, obtuse to rounded. Androecium with fertile
stamens inserted 0.7–1.3 mm from the base of the corolla; shorter ones 2–3 mm long; longer ones 2.5–4 mm long;
anthers thecae 0.6–0.8 mm long, obovate to elliptic, subexserted; connective extending 0.1–0.3 mm beyond anther
attachment; staminode 0.9–1.2 mm long, glabrescent, with long and short stipitate trichomes. Gynoecium 3.2–5.1
mm long; ovary 0.7–1.3 mm long, 0.5–1 mm wide, conical, velutinous; style 2–3.9 mm long, tomentose at the
base. Fruit a linear flattened capsule, 10.5–25 cm long, 0.4–0.9 cm wide, coriaceous to woody, granular throughout
(sometimes smooth near the midvein), without lenticels to densely lenticeled, villous to puberulent or glabrescent,
with simple and peltate trichomes; central ridge single, slightly or not prominent; margins slightly raised
(unwinged), ca. 0.1 cm wide. Seeds body 0.7–1.5 cm long, 0.3–0.9 cm wide; wings (0.2–)0.4–1.2 cm long.
Phenology:—Flowers throughout the year and produces fruits from March to August.
Distribution and habitat:Occurs in moist broadleaf forests from Bolivia (Beni, Cochabamba, La Paz,
Pando and Santa Cruz), Brazil (Acre, Amazonas, Mato Grosso, Pará and Rondônia), Colombia (Antioquia, Chocó
and Meta), Dominican Republic (El Seybo, Monte Cristi and Samaná), Ecuador (Morona-Santiago, Napo and
Zamora-Chinchipe), Guiana (Upper-Essequibo), Peru (Amazonas, Cusco, Huánuco, Junín, Loreto, Madre de Dios,
Pasco, Puno, San Martín and Ucayali), Porto Rico and Venezuela (Amazonas) (Fig. 18).
FIGURE 18. Distribution of Tyna nth us pol yan thu s.
Additional specimens examined:—Sin loc.. October 1831, E.F. Poeppig 2651 (LE, P). BOLIVIA. Beni:
Moxos, 66 km de San Ignacio, San Borja, 200 m, 20 February 1995, M. Moraes & E. Oviedo 2074 (MO). Vaca
Diéz, 11–15 km NW of Guayaramerin on road to Cachuela Esperanza, 12 February 1978, W. R. A nd er so n 12 047
(MO, NY, US); Riberalta, 160 m, 29 June 1992, A.H. Gentry et al. 77563 (MO). Cochabamba: Carrasco, Entre
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Ríos, Dist. Tres, Sector Coop. Litoral, 280 m, 12 July 2005, O. Colque & L. Mendoza 262 (SPF); 6 February 2006,
O. Colque & L. Mendoza 410 (SPF). Cercado, The road from Ivirgarsama, approximately 2 km towards Puerto
Villarroel, 200 m, 24 February 1996, N. Ritter 2996 (MO). Chapare, Villa Tunari, Hotel El Puente, 300 m, 21
January 1996, J.R.I. Wood 10423 (K). La Paz: Sin loc., 1–22 July 1939, B.A. Krukoff 10456 (K). Abel Iturralde,
Tumupasa, 540 m, 10 January 1902, R.S. Williams 508 (K, NY); Siete Cielos, Río Manupare, 180 m, 4 June 1987,
J.C. Solomon 16919 (MO). Franz Tamayo, Parque Nacional Madidi, río Quendeque, 300 m, 7 February 2002, A.
Fuentes et al. 3753 (MO); Ibid., NW de Apolo, senda Azariamas–San Fermin, Parcela temporal (0.1 ha), 1326 m,
27 May 2006, I. Loza et al. 73 (SPF); 27 May 2006, I. Loza et al. 78 (SPF). Nor Yungas, Below San Pedro,
Yolosa–Caranavi road, valley of Rio Coroico, 760–950 m, 15 January 1984, A.H. Gentry et al. 44262 (M, MBM,
MO, NY); Corocoro, 12 km NE of Caranavi, 1400–1500 m, 16 January 1984, A.H. Gentry et al. 44315 (MO, NY);
16 January 1984, A.H. Gentry et al. 44350 (MO); 1 km SW of Yolosa on road to Chuspipata, Ford of Río Cedro,
1400 m, 22 February 1984, J.C. Solomon & J. Kuijt 11594 (F photo, MO); 5.5 km below Coroico (towards Yolosa)
(1.4 km above Yolosa), 1400 m, 16 May 1985, J.C. Solomon 13733 (MBM, MO, NY); 3.8 km below Coroico and
Yolosa, 1500 m, 25 September 1986, J.C. Solomon 15665 (MO). Sud Yungas, Basin of Rio Bopi, Asunta (near
Evenay), 690–750 m, 27–31 July 1939, B.A. Krukoff 10695 (F, G, K, MO, NY, US); Valle del río Unduavi, entre
Santa Rosa y Machacamarca expo. N-0, 1400 m, 4 February 1988, R. Seidel & E. Richter 1207 (MO). Pando:
Abuná, Nuevo Mundo, 180 m, 29 June 1992, A.H. Gentry et al. 77573 (MO). Nicolás Suárez, Km 15 between
Cobija and Porvenir, 300 m, s.d., M. Cardenas 4167 (US); En la zona de Campoana, junto a la barraca San José,
290 m, 14 January 1983, F. Casas & Susanna 8264 (G, MO, NY). Santa Cruz: Im Gebüsch des Waldes am
Westufer des Rio Pirai, unweit Santa Cruz, January 1911, T. Herzog 1460 (G). Andrés Ibáñez, 2 km W of center of
La Belgica, 360 m, 24 January 1987, M. Nee 33781 (G, MO, NY); 4 km S of Pedro Lorenzo on old road from Santa
Cruz to Abapó, 525 m, 21 January 1998, M. Nee 48058 (MO, NY); Along "old" highway to Cochabamba, vic.
Turnoff of highway south to Abapó, 13 km SW of center of Santa Cruz, 450 m, 29 July 2003, M. Nee 52391a
(NY); 6.4 km (by dirt roads) SE of Pedro Lorenzo on way to Peji, 4.5 km SW of Peji, 530 m, 29 June 2004, M. Nee
52633 (MO, NY); Camino entre Warnes y La Bélgica, 353 m, 18 February 2006, D. Villarroel et al. 347 (NY, SPF);
Porongo, Localidad del Monumento Natural Espejillos, Precipicio del mirador, 1200 m, 16 January 2008, I. Linneo
& D. Galindo 1326 (SPF). Cercado, Bosques del Piray, 450 m, 4 February 1917, J. Steinbach 3224 (BM, F photo,
K); 20 January 1921, J. Steinbach 5253 (F, G, MO, NY). Cordillera, 2.5 km (by road) W of railroad and 3.5 km (by
road) W of the Santa Cruz–Abapó highway and San Joaquín, along gravel road to "Pozo PCHX1001", the turnoff
5.5 km S of Basilio, 570 m, 24 May 2005, M. Nee 53129 (MO, NY, SPF). Ichilo, Santa Cruz 130 km NW linea
recta, puente Rio Yapacaní 35 kms hacia Puerto Grether, 250 m, 18 March 1981, S.G. Beck 6557 (M, MO); Vicinity
of old buildings for highway construction, ca. 2 km W of Villa San Germán on highway from Buena Vista to Río
Ichilo, 275 m, 22 February 1998, M. Nee 48385 (MBM, MO, NY); 3.5 km SW of turnoff at Villa San Isidro from
new highway from Buena Vista to Río Ichilo, 360 m, 1 March 1998, M. Nee 48546 (MO, NY); Buena Vista, Orillas
del camino entre Buena Vista y el Cairo, 320 m, 2 February 1990, I.G. Vargas C. 377 (MO, NY); Ignacio Warnes, E
side of the village of Colonia Okinawa 1, 250 m, 28 January 1987, M. Nee 33837 (MO, NY). José Miguel de
Velasco, Parque Nacional Noel Kempff Mercado, 14 km SE del Estancia Flor de Oro, cerca Rio Itenez, 260 m, 9
March 1992, A. Perry 686 (MO); Ibid., Comunidad Florida, 200 m, 2 July 1993, M. Saldias et al. 2928 (MO); Ibid.,
Lago Caiman, 540 m, 19 January 1997, M. Garvizu et al. 336 (G, MO); Ibid., Campamento Los Fierros, 200 m, 18
March 1997, L. Sánchez et al. 402 (G, MO); Carretera al Chore, Entre Arroyo Las Londras y Arroyo El Tigre, 150
m, 24 July 1996, M. Peña-Chocarro & L. Arroyo 167 (G, MO); 25 km al norte del aseradero Moira yendo al Chore,
Bosque de lianas del "Proyecto Bejucal" de USZ, 160 m, 26 July 1996, A. Jardim et al. 3189 (MO); Camino de
Florida hacia las Mechitas, 450 m, 24 January 1997, J. Guillen et al. 297 (G, MO). Ñuflo de Chávez, Perseverancia,
NW of Santa Cruz on Río Negro, Transect 3, 100 m, 11 May 1991, A.H. Gentry & M. Peña 73697 (MO); Near San
Julian (S), 274 m, 27 February 2003, L. Rico et al. 1406b (K, MO). Sara, Área natural de manejo integrado
Amboró, camino a San Isidro, pasando el puente del río Moyle, 278 m, 20 February 2006, D. Villarroel et al. 367
(NY, SPF). BRAZIL. Acre: Jordão, Trails and roads 2–3 km SE of Foz de Jordão, 250–275 m, 7 February 2009,
F. A . M i c he l a n ge l i e t al . 1 35 6 (SPF). Acrelândia, BR-364, km 87, Ramal do Projeto 2, 13 February 2000, I.S.
Rivero et al. 309 (SPF). Bujari, Basin of Rio Purus, Riozinho do Andirá (tributary of Rio Acre), Seringal Andirá,
24 March 1995, D.C. Daly et al. 8457 (MO, NY, UFACPZ); Riozinho do Andirá, Colocação Curitiba, 8 June 1995,
A.R.S. Oliveira et al. 540 (SPF, UFACPZ); Rio Antimari, Floresta Estadual do Antimari, right bank, Colocação Pé
da Terra, 11 March 1997, D.C. Daly et al. 9474 (SPF, UFACPZ). Cruzeiro do Sul, Perto do entroncamento com o
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TAXONOMIC REVISION OF TYNANTHUS (BIGNONIEAE)
ramal Pentecoste, 26 April 2001, L.G. Lohmann et al. 370 (MO, NY). Marechal Thaumaturgo, Reserva Extrativista
do Alto Jurua, Rio Juruá, S of confluence with Rio Acuriá and N of São João do Breu, Colocação Tapaúna, 31
March 1993, D.C. Daly et al. 7653 (INPA, NY, UFACPZ); Ibid., Fazenda Paraguay, 3 April 1993, D.C. Daly et al.
7728 (INPA, NY, UFACPZ); Ibid., Foz do Bajé, Boa Vista da União, 30 April 2001, L.G. Lohmann et al. 413 (MO,
NY); Ibid., Foz do Bajé, Boa Vista da União, Colocação Horizonte, 1 May 2001, L.G. Lohmann et al. 421 (MO,
NY); Ibid., Margem direita do Rio Arara, 6 May 2001, L.G. Lohmann et al. 476 (MO, NY). Plácido de Castro,
Porto Edith, arredores da casa do Sr. João, Margem esquerda do Rio Abunã, 14 May 2001, L.G. Lohmann & E.C.
Oliveira 505 (MO, NY). Porto Acre, Reserva Florestal do Humaita, Beira do Rio Acre, 21 March 1995, C.
Figueiredo et al. 741 (SPF, UFACPZ). Rio Branco, Parque Zoobotânico, Estrada Dias Martins, próximo à entrada
do Herbário, s.d., G. C l a r o s & R . E h r i c h 3 1 5 (UFACPZ); Ibid., 3 March 1993, C.S. Figueiredo & H.B.N. Borges
100 (UFACPZ); Ibid., inside main gate, 25 March 2002, D.C. Daly 11565 (NY); Ibid., estrada de acesso ao
herbário, 160 m, 5 March 2012, M.C. Medeiros et al. 37 (SPF); Km 1 ao km 23 da Rodovia Rio Branco–Porto
Ve lh o, 2 1 F eb r ua r y 1 97 8 , J.U. Santos et al. 69 (MG, MO, US); Rio Branco–Porto Velho Highway, between km
22–37, 8 February 1979, B.W. Albuquerque et al. 1345 (MO, NY, US). Santa Rosa, Rio Purus, left bank, Seringal
Refúgio, S of Igarapé Extrema de Baixo, 25 March 1999, D.C. Daly et al. 10083 (SPF, UFACPZ). Sena Madureira,
BR-364, entre km 245–246, Fazenda Liberdade, 140–150 m, 6 February 2002, M. Alves et al. 2734 (SPF); Basin of
Rio Iaco (tributary of Rio Purus), Fazenda São Jorge I, property of Acre Brasil Verde, timber concession of
Laminados Triunfo Ltda, 107 km NW of Rio Branco on BR-364, then ca. 22 km E on Toco Preto access road, 200
m, 7 July 2008, D.C. Daly et al. 13228 (RB). Xapuri, Fazenda Bomfim, 18 March 1995, D.C. Daly et al. 8365
(MO, NY, UFACPZ); 19 March 1995, D.C. Daly et al. 8386 (MO, NY, UFACPZ); Margem direita do Rio Xapuri,
18 May 2001, L.G. Lohmann & E.C. Oliveira 567 (NY); Estrada para Brasiléia (BR-317) km 243, ramal da
Cachoeira, a aproximadamente 8 km da BR, em frente à fazenda Bom Jesus, 200 m, 9 March 2012, M.C. Medeiros
et al. 40 (SPF). Amazonas: Along Rio Castanho, tributary of Rio Padauiri, upper Rio Negro Basin, 100–140 m,
16–24 February 1946, F. C a r d o n a 1 4 1 9 (US). Esperança, Ad ostium fluminis Javary, 11 February? 1942, A. Ducke
866 (F, IAN, MG, MO, NY, R, RB, US). Lábrea, Rio Curuquetê, Cachoeira Republica, 24 July 1971, G.T. Prance et
al. 14559 (INPA, K, MG, MO, NY, R, US). Manaus, Road to Rio Negro from km 12 of Estrado do Aleixo, vicinity
of Manaus, 14 December 1974, A.H. Gentry 13303 (INPA, MG). Parintins?, 1 April 1946, J.M. Pires & G.A. Black
1211 (K, IAN). Presidente Figueiredo, Estrada Manaus–Caracaraí, Reserva Biológica do INPA, 1978, A. Anderson
s.n. (INPA 142420). Mato Grosso: Salto do Sepotuba, March 1909, F. C . H o eh n e 1 5 6 8 ? (R); March 1909, F.C.
Hoehne 1582 (R); March 1909, F.C. Hoehne 1583 (R). Novo Mundo, Reserva Particular do Patrimônio Natural
Lote Cristalino, Trilha do Teles Pires, 172 m, 7 May 2007, D. Sasaki et al. 1625 (K); Parque Estadual Cristalino,
Cachoeira do Escondido, entrada pela Fazenda AJJ, 28 January 2008, D. Zappi et al. 900 (K). Tapirapoan, January
1914, F.C. Hoehne 5852 (R); January 1914, F.C. Hoehne 5853 (R). Pará: Rio Branco de Óbidos, Castanhal grande,
11 December 1913, A. Ducke s.n. (MG 15122); 25 January 1918, A. Ducke s.n. (MG 16943). Roadside on BR 163,
Cuiabá–Santarém road, km 879, 310 m, 14 February 1977, J.H. Kirkbride Jr. & E. Lleras 2787 (F photo, INPA, K,
MG, MO, NY, RB, US). Altamira, Xingu River, Assurini Indians, 15 June 1986, W. Balée 2597 (NY). Faro, 19
August 1907, A. Ducke s.n. (K, MG 8392); 30 January 1910, A. Ducke s.n. (INPA 11254, K, MG 10565); 11 May
1911, A. Ducke s.n. (MG 11647). Itaituba, Rio Tapajós, s.d., A. Ducke s.n. (MO 2305625, R 23820, RB barcode
58413). Jacundá, Rio Tocantins, 14 May 1951, R.L. Fróes 27092 (IAN, K, RB). Marabá, Serra dos Carajás, estrada
do estéril sul, às proximidades da Barragem, 6 February 1985, O.C. Nascimento & R.P. Bahia 1189 (MG). Monte
Alegre, Colonia Itauajury, 6 March 1923, A. Ducke s.n. (R 23821, RB barcode 58425). Oriximiná, Rio Trombetas
ao norte, ao longo do rio próximo à Mineração Rio Norte, 22 July 1980, C.A. Cid et al. 1710 (F, G, INPA, MG, MO,
NY, US); Porto Trombetas, Mineração Rio do Norte, Entrada do Alter do chão, 20 November 1990, O.H. Knowles
1620 (INPA); Ibid., 1991, Evandro & Knowles 424 (INPA); Comunidade Pancada, 22 June 2006, D.R. Oliveira &
Augusto 98 (INPA). Parauapebas, Serra dos Carajás, Vila de N-5, final da rua Angelim, 29 March 1989, J.A.A.
Bastos 161 (MG). Pau D'Arco, Marajoara, s.d., J. Grogan 51 (IAN); 16 February 1997, J. Grogan 50 (IAN, INPA).
Santarém, Serra de Pequiatuba, 28 March 1924, J.G. Kuhlmann 1816 (MO, R, RB); Estrada Santarém–Cuiabá, km
67, Reserva florestal do IBDF, 6 March 1979, M.R. Cordeiro et al. 1610 (IAN); FLONA Tapajós, BR-163, km 83,
estrada de acesso à torre do projeto LBA, 150 m, 21 November 2011, M.C. Medeiros & T. André 36 (SPF).
Tucuruí, BR-423, 15 March 1981, U.N. Maciel & C.S. Rosário 552 (MG). Rondônia: Estrada Porto
Ve lh o – Cu i ab á , B R - 36 4 , k m 2 90 , ma r g em e sq u er d a d a es t r ad a , 1 3 Fe b r ua r y 1 9 83 , C.A.A. Freitas et al. 98 (INPA,
MO). Porto Velho, Campo da sub-estação, 18 November 1949, N.T. Silva 358 (IAN, INPA, K); Guajará-
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Phytotaxa 216 (1) © 2015 Magnolia Press
Mirim–Abunã, trecho entre o km 12 ao km 36, 1 February 1983, L. Carreira et al. 414 (IAN, INPA, MG, NY).
Presidente Médici, Picadão que confina a 7a. e 8a. linha, margem esquerda da BR 429, 24 March 1986, N.A. Rosa
et al. 4984 (MG, MO). COLOMBIA. Antioquia: Caucasia, Hacienda "La Candelaria", Universidad de Antioquia,
150 m, 5 September 2000, R. Fonnegra et al. 7157 (MO). Chocó: Upper Río Truandó, La Teresita (INDERENA
Camp.), 100–200 m, 19 January 1974, A.H. Gentry 9383 (MO). Meta: Villavicencio, 450 m, 1–2 September 1917,
F.W. Pennell 1602 (NY). DOMINICAN REBUBLIC. El Seybo: El Jovero, wooded slope at La Pocilga, 150 m, 27
July 1930, E.L. Ekman H.15782 (K, NY, US). Monte Cristi: Sabaneta, La Cidra, In vicinity of Laguna, 5–600 m, 9
November 1930, E.J. Valeur 532 (K, MO, NY, US); Ibid., 640 m, 7 April 1955, J.J. Jimenez et al. 2909 (K, US).
Samaná: Vicinity of Laguna, Samaná Peninsula, chiefly on the Pilón de Azúcar, 100–500 m, 19 December 1920,
W.L . Ab bo tt 33 0 (US); Sabana de la Mar, in Loma del Bejuco-clavo, 7 July 1930, E.L. Ekman H.15590 (US); La
Laguna, Península de Samaná, 75 m, 5 August 1930, E.L. Ekman H.15853 (F, MO). ECUADOR. Morona-
Santiago: Basin of Rio Morona, Rio Mangosiza, Mission of Miazal, 45 km ESSE of Sucua (by air), at edge of
cultivated field at Chumbee's house, 300 m, 19 February 1990, C.F. Limbach & M.D. 125 (MO). El Centro Shuar
Pampants, Rio Kankaim (Cangaime), 300 m, 13 September 1985, A. Warush RBAE99 (MO, NY). El Centro Shuar
Kankaim (Cangaimine), Rio Kankaim (Cangaime), 20 km WNW del Taisha, 500 m, 15 October 1985, D. Shiki
RBAE216 (NY); 15 October 1985, D. Shiki RBAE218 (MO, NY). Morona, Cordillera de Cutucú, Centro Shuar
Uunsuants/Transkutuku, Bosque intervenido junto al Río Mainpaimi, 600 m, 18 January 2002, W. P al aci os e t al .
15532 (MO). Napo: Road from Lago Agrio Airport, 250 m, 11 February 1974, A.H. Gentry 9836 (MO). 9–11 km
S of Coca on road to Auca oil field, 5 November 1974, A.H. Gentry 12491 (MO). 43–47 km S of Coca, end of
Aucas oil field near Río Tiputini, 300 m, 5 November 1974, A.H. Gentry 12512 (MO). 4 km sur de Puerto Napo em
el Rio Napo, 500 m, 4 August 1984, C. Dodson et al. 14934 (MO). Estación Experimental INIAP-Payamino, 5 km
NE de Coca, 250 m, 16–26 February 1986, D. Neill 7151 (MO, NY). Reserva Etnica Huaorani, Carretera y
oleoducto de Maxus, km 119–120, Carretera a Plataforma GINTA, 235 m, March 1995, M. Aulestia et al. 3586
(SPF). Yasuni Forest Reserve, road from PUCE Scientific Station to end of road towards Waoroni Territory,
240–310 m, 29 June 1995, P. A c e v e do - R d g z . & J . A . C e d e ñ o 7 5 72 (MO, P, US). Tena, Estación Biológica Jatun
Sacha, Río Napo, 8 km al este de Misahualli, Parcela Permanente 02, 400 m, 14 December 1989, W. Palacios 4771
(G, K, MO, NY). Zamora-Chinchipe: Nangaritza, Miazi, along Rio Nangaritza, Transect 3, 850 m, 28 July 1993,
A.H. Gentry 80558 (MO). GUIANA. Upper Essequibo: Rewa River, near Camp 2 at foot of Spider Mountains,
200–250 m, 22 September 1999, M.J. Jansen-Jacobs et al. 6073 (F, K, MO, NY, P, US). PERU. Amazonas:
Bagua, Quebrada shimpunts alto rio Cenepa, Monte al lado shimpunts, 800–1100 ft, 22 February 1973, E. Ancuash
41 (F photo, MO, NY); Quebrada Huampami, Rio Cenepa, Monte orilla de Huampami, 600 ft, 29 May 1973, R.
Kayap 829 (F, MO); Marañon valley, between Oracusa and Sta. Maria del Nieva, 240 m, 11 February 1978, D.C.
Was sh ausen & F. E ncar nación 884 (G, K, MO, US); 2–3 km SW of Chiriaco on road to Bagua Chica, Marañon
Va ll e y, va l le y of Rí o C h ir i ac o , 3 00 –3 3 0 m , 1 7 Ju n e 1 97 8 , A.H. Gentry et al. 23123 (MO); Imaza, Comunidad
Aguaruna Yamayakat, 240 m, 23 March 1995, C. Diaz et al. 7597 (F, MO); Ibid., 320 m, 13 March 1996, N.
Jaramillo et al. 1374 (F, MO); Ibid., Quebrada Kusú, transecto 2 × 500 m, 380 m, 1 November 1996, R. Vásquez et
al. 21375 (MO); 5 November 1996, R. Vásquez et al. 21497 (MO); 9 November 1996, R. Vásquez et al. 21706
(MO); 9 November 1996, R. Vásquez et al. 21772 (MO); Ibid., 600 m, 6 June 1997, R. Vásquez et al. 23937 (F,
MO, NY). Condorcanqui, Huambisa, Valle del Rio Santiago, approx. 65 km N de Pinglo, Quebrada Caterpiza, 200
m, 10 March 1980, V. Huashikat 2218 (MO); El Cenepa, Comunidad Aguaruna Pagki-Suwa, Río Cenepa, 289 m,
21 January 1997, R. Vásquez et al. 22079 (F, MO). Cusco: La Convención, Rio Manguriari (Manguyari), Alto
Urubamba, upstream to Rio Manguriari, 750 m, 2 February 1991, P. N ú ñ e z & G. O r t i z 1 27 5 8 (MO); 2 February
1991, P. Núñez & G. Ortiz 12837 (MO); 2 February 1991, P. Nuñez & G. Ortiz 12845 (MO); Echarate, San Martin-
3 Well Site, 400 m, 10 March 1997, P. N u ñ e z e t a l . 1 9 7 2 0 (NY, US). Quispicanchi, Hills around Río Araza between
Pande Azucar and Quince Mil Airport, 292 km from Cusco, 643 m, 10 August 1991, P. N u ñ e z V. 1 3 9 0 5 (MO); 10
August 1991, P. Núñez 14080 (MO); Camanti, 254 km from Cusco road to Maldonado, 15 Quince area, 643 m, 18
February 1991, P. Núñez 13006 (MO). Huánuco: Leoncio Prado, Vicinity of Tingo María, East of Río Huallaga,
670 m, 9 March 1962, J. Schunke Vigo 5813 (F, K, MO, US); Hills east of Tingo Maria, 5 October 1972, T.B . C roa t
21121 (MO); Across Río Huallaga from Tingo Maria, 650 m, 17 January 1976, A.H. Gentry et al. 15894 (F, MO,
NY); Rupa Rupa, Tingo María, Limestone hills opposite airport, 700–780 m, 9 December 1981, T. Plowman et al.
112 76 (F, K); Tingo Maria, 670 m, 22 August 1959, F. Wo y t ko w s ki 5 3 91 (K, MO). Marañón, Valley of Rio
Huallaga, 110–140 km N of Tingo Maria, near San Martin border, 550 m, 4 February 1984, A.H. Gentry & D.N.
Phytotaxa 216 (1) © 2015 Magnolia Press
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TAXONOMIC REVISION OF TYNANTHUS (BIGNONIEAE)
Smith 44929 (MBM, MO, NY). Pampahuasi, January 1830, E.F. Poeppig 1617 (G). Junín: La Merced, 700 m, 20
May–4 June 1929, E.P. Killip & A.C. Smith 23670 (US). Loreto: Tarapoto, November 1902, E. Ule 6577 (G, K,
MG). Alto Amazonas, Fortaleza, near Yurimaguas, 140 m, December? 1932, G. Klug 2778 (BM, F, G, K, MO, NY,
US); Old Andoas, Río Pastaza, 190 m, 26 December 1985, W.H . Le wi s & M .C . Gn er re 1 0 37 2 (MO); Washintsa and
vicinity, Río Huasaga, Achual Jívaro, 185 m, 16–26 June 1986, W.H. Le wi s et al. 111 25 (MO); Puranchim, Río
Sinchiyacu, 200 m, 21–27 November 1986, W.H . L ew is e t al . 1184 9 (MO); Petro Estación Morona, río Morona,
160 m, 22 March 1987, W.H . L ew is et al . 1293 4 (MO); Andoas, Río Pastaza near Ecuador border, near airport, 230
m, 16 November 1979, A.H. Gentry & C. Díaz S. 28135 (F, MO); Ibid., 210 m, 15 August 1980, A.H. Gentry et al.
29814 (F, MO); Balsapuerto, 220 m, January 1933, G. K l u g 2 8 3 6 (BM, F, G, K, MO, NY, US). Coronel Portillo,
IVITA, km 59 from Pucallpa toward Tingo Maria, 250 m, 16 January 1976, A.H. Gentry & J. Revilla 15888 (MO).
Datem del Marañón, Tierra Blanca, on Rio Morona?, s.d., G. T e s s m a nn 4 9 1 4 (G, NY). Loreto, Nueva Jerusalem and
vicinity, Río Macusari, 220–300 m, 29 December 1985–3 January 1986, W.H . Le wi s et a l. 1 04 21 (MO); Ibid.,
10–11 June 1986, W.H. Lewi s et al . 1 09 87 (MO); Tigre, Río Corriente, Teniente López, Caserio Jíbaros, Bosque 1o,
26 November 1979, F. Ayala et al. 2428 (MO); Pampa Hermosa and vicinity, Río Corrientes, 1 km al S of junction
with Río Macusari, 160 m, 3–20 December 1985, W.H. Lewis et al. 10184 (MO); Nueva Nazaret, Río Morona, 160
m, 3 December 1986, W.H. L ewi s e t al . 1 2445 (MO). Maynas, Rio Zumun, affluent du rio Yahuas-Yacu, affluent du
rio Ampi-Yacu, affluent de l'Amazone à Pebas, commune de Colonia, territoire des Indiens Bora, 14 June 1978, S.
Barrier 1095 (F, G, NY, P); Caseria Alianza, Río Tamshiyacu, trail toward Río Manití, 140 m, 1 August 1980, A.H.
Gentry et al. 29319 (MO); 1 August 1980, A.H. Gentry et al. 29321 (MO); Yanamono, Explorama Tourist Camp,
Río Amazonas halfway between Indiana and mouth of Río Napo, 130 m, 10 July 1983, A.H. Gentry et al. 42873
(MO); Alto Nanay, Near Santa Maria de Nanay, 24 February 1968, D.R. Simpson & J. Schunke Vigo 702 (G, NY,
US); Iquitos, Manacamiri, Rio Nanay, 130 m, 5 March 1995, F. A ya l a 68 6 7 (M); Quistococha, Ca. 10 km of
Iquitos, Arboretum of Universidad de Amazonas, 15 November 1974, A.H. Gentry & F. Ayala 12689 (F, MO).
Madre de Dios: Manu, Parque Nacional del Manu, Rio Manu, Cocha Cashu Station, 400 m, 3 April 1977, B. Bell
& R.B. Foster 6262 (F photo); Ibid., 380 m, 17 October 1979, A.H. Gentry et al. 26862 (F, MO); Pantiacolla,
Serrania across Río Alto Madre de Dios from Shintuya, 450–650 m, 28 October 1979, A.H. Gentry et al. 27282 (F,
MO). Tambopata, Ca. 5 km from Puerto Maldonado near Río Tambopata, 200 m, 24 January 1976, A.H. Gentry &
J. Revilla 16318 (F, MO); Road from Puerto Maldonado to Tambopata, 0–4 km from Puerto Maldonado, 220 m, 25
January 1976, A.H. Gentry & J. Revilla 16354 (F, INPA, MO, NY); Ca. 20 km W of Puerto Maldonado, on road to
Quince Mil, 250 m, 23 April 1977, A.H. Gentry et al. 19698 (F, MO); Outskirts of Puerto Maldonado, 210 m, 27
February 1981, A.H. Gentry & K. Young 31757 (F, G, MO); Cusco Amazónico, Río Madre de Dios below Puerto
Maldonado, 200 m, 20 February 1989, A.H. Gentry & P. Nuñez 66032 (MO); Lago Sandoval, 13 km NE of Puerto
Maldonado, lake edge, trails, 200 m, 25 July 1989, P. Nú ñe z 11 19 9 (MO); Cuzco Amazónico, trail to Lago
Sandoval across Río Madre de Dios, ca. 12 km E of Puerto Maldonado, 200 m, 21 February 1990, A.H. Gentry & P.
Núñez 69375 (MO); Río Heath, Peru/Bolivia border, 200 m, 2 March 1990, A.H. Gentry & P. Núñez 69786 (MO,
NY); 2 March 1990, A.H. Gentry & P. Núñez 69793 (MO, NY); Puerto Maldonado, 650 ft, 21 March 1944, R.J.
Seibert 1899 (MO, US). Pasco: Oxapampa, Palcazu valley, near the confluence of Rio Palcazu and Rio Iscosacin,
Juan Frantzen property, 300 m, 23 April 1983, D.N. Smith 3890 (MO). Puno: Carabaya, Ridge between Río
Candamo and Río Guacamayo, 400–600 m, 22 May 1992, A.H. Gentry et al. 76975 (MO, NY). San Martín:
Yurimaguas a Huallaga, 1924, G. Te s s m a n n 5 5 1 3 (G). Tingo Maria, Near 21810, 625–1100 m, 30 October 1949–19
February 1950, H.A. Allard 21811 (US). Mariscal Cáceres, Juanjui, Alto Rio Huallaga, 400–800 m, December
1935, G. Klug 4160 (BM, F, K, MO, NY, US); December 1935, G. Klug 4194 (BM, F, K, MO, NY, US); Tocache
Nuevo, Camino al pueblo viejo de Tocache, 18 January 1970, J. Schunke Vigo 3732 (F, NY, US); Aeropuerto de
Tocache, 400 m, 26 January 1970, J. Schunke Vigo 3771 (F, G, NY, US); Al Nor-Oeste del vivero del Instituto
Agropecuario de Tocache, 18 April 1970, J. Schunke Vigo 3943 A (F photo); Almendras, camino a pueblo Viejo,
400 m, 5 April 1975, J. Schunke Vigo 8216 (MO, NY); Quebrada de Cachiyacu, afluente de la quebrada de
Huaquisha, 500–600 m, 17 May 1975, J. Schunke Vigo 8481 (MO, NY); Quebrada de Ishichimi, cerca a Tocache,
400 m, 11 March 1978, J. Schunke Vigo 10004 (F, MO, NY); Farm of Don Roberto Aguillar, 450 m, 1 July 1978, T.
Plowman & J. Schunke Vigo 7509 (F, MO); Cerro de Sin Sin, cerca a Bambamarca, 600–630 m, 20 January 1979,
J. Schunke Vigo 10729 (MO); Trail up Río Huallaga Valley toward Limón, 500 m, 11 March 1979, A.H. Gentry et
al. 25519 (F, MO, NY); Río de la Plata, cerca a la chacra del Sr. Esteban Arévalo, al borde de la quebrada, 650 m,
14 September 1980, J. Schunke Vigo 12280 (F, K, MBM, MO, NY, RB); Río de la Plata, cerca a la Chacra del Sr.
MEDEIROS & LOHMANN
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Phytotaxa 216 (1) © 2015 Magnolia Press
Esteban Arévalo, 650 m, 15 September 1980, J. Schunke Vigo 12298 (MO). Fundo Jeroglífico, propriedad del Sr.
Luis Ludeña, al Sur de Tocache, 400 m, 20 July 1988, J. Schunke Vigo 14266 (MO). Rioja, Along road
Yorongos–La Florida near Rioja, 1000 m, 31 March 2001, H. van der Werff et al. 16551 (F, MO, NY). San Martin,
Chazuta, Río Huallaga, 260 m, April 1935, G. Klug 4065 (BM, F, K, MO, NY, US); Ibid., Llucanayacu, 300 m, 3
November 2004, J.S. Biset 43 (SPF); Ibid., 400 m, 23 May 2005, J.S. Biset & J.C. de la Cruz 336 (SPF); Ibid., 24
May 2005, J.S. Biset & J.C. de la Cruz 370 (SPF); Ibid., Llucanayacu, 600 m, 26 May 2005, J.S. Biset & J.C. de la
Cruz 453 (SPF). West side of Río Huallaga, West of Shapaja 2–8 km on trail to Tarapoto in the Cerros de Estoraqui,
1300 ft, 4–7 August 1937, C.M. Belshaw 3217 (K, MO). Ucayali: Coronel Portillo, Bosque Nacional Alexander
von Humboldt, km 86 Pucallpa–Tingo Maria road, 300 m, 27 March 1977, A.H. Gentry et al. 18697 (MO); 27
March 1977, A.H. Gentry et al. 18739 (MO); Ibid., 270 m, February 1978, C. Froehner 182 (MO, NY); Ibid., 250
m, 8 March 1979, A.H. Gentry et al. 25475 (INPA, MO, NY); Ibid., 270 m, 9 February 1981, A.H. Gentry et al.
31133 (G, MO, NY); Aguaytia, Woods to north of house, Don Diogenes del Aguila, 30 June 1960, M.E. Mathias &
D. Taylor 5062 (MO, US); Campoverde, Km 36, CF Basadre, 250–270 m, 27 October 1988, F. C h a v ez 3 (MO).
VENEZUELA. Amazonas: Alto Orinoco, Raudal de los Guaharibos, 24 July 1951, L. Croizat 335 (F, NY). 22–23
km N of Samariapo on road to Puerto Ayacucho, 100 m, 29 June 1975, A.H. Gentry & P. Berry 14572 (MO, US).
Taxonomic notes:Tynanthus villosus was described as a new species based on the “shaggy indumentum”,
“narrow inflorescences” and “persistent prophylls” (Gentry 1976: 60). Nevertheless, analyses of the collections of
T. polyanthus and T. villosus throughout their distribution ranges indicated a continuum in the patterns of variation
in the indumentum and inflorescence morphology, with vegetative portions from both species varying from villous
to pubescent or glabrescent, and the inflorescences varying from large, compound and multi-flowered to small,
simple or bearing only a few flowers. A careful analysis of specimens collected at different developmental stages
(e.g., vegetative, flowering and fruiting) indicated that the prophylls of the axillary buds are generally lost in older
specimens. Apart from the morphological similarities between these taxa, T. villosus appeared nested within T.
polyanthus in the molecular phylogeny of the genus (Medeiros & Lohmann 2015). Our morphological and
molecular phylogenetic observations thus led us to accept a more broadly circumscribed T. polyanthus and to
synonymize T. villosus in T. polyanthu s. As circumscribed here, T. polyanthus is characterized by the
conspicuously tetragonal and angular young branchlets, simple tendrils, foliaceous prophylls of the axillary buds,
lax inflorescences, short corollas (0.4–0.8 cm long), and unwinged fruits. Tynanthus polyant hus is sister to T.
cognatus (Medeiros & Lohmann 2015), with which it shares some morphological features (see taxonomic notes
under T. cognatus).
Nomenclatural notes:—In the protologue of Schizopsis polyantha, Bureau cited a specimen deposited at G.
However, no duplicates of Spruce 4895 were located at G during a visit in 2006 during which time materials were
analyzed for the recently published Bignonieae synopsis (Lohmann & Taylor 2014). Therefore, Lohmann, in
Lohmann & Taylor (2014), designated the specimen Spruce 4895 deposited at P, where most of Bureau’s types are
located, as the lectotype of T. polyanthus. In 2013, however, two duplicates of the collection Spruce 4895 were
located at the G herbarium, indicating that the G collection cited in the protologue of S. polyantha was not lost or
destroyed as previously thought. Despite that, we here follow the lectotypification proposed in Lohmann & Taylor
(2014).
As far as T. m yrianthus is concerned, Bureau & Schumann (1896) did not clearly indicate in the protologue
which of the duplicates of the collection Poeppig 2388 was the lectotype. We here select the duplicate deposited at
W, which has be en broadl y phot ographed and distr ibuted as the hol otype of T. myrianth us as its lectotype.
Similarly, the best quality material of the collection C.A.M. Lindman A2967 from S is here selected as a lectotype
for T. lindmanii.
A neotypification of T. caryophylleus was necessary given that all of Bello’s types kept at B (Stafleu & Cowan
1976: 170) were subsequently destroyed during World War II (Hiepko 1987). Since no duplicates of those
specimens were distributed to any other collection, a specimen of T. caryophylleus from Porto Rico (where the
original type was collected), identified by Alain (author of the combination of T. caryophylleus) is here selected as
the neotype.
Phytotaxa 216 (1) © 2015 Magnolia Press
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TAXONOMIC REVISION OF TYNANTHUS (BIGNONIEAE)
12. Tynanthus pubescens Gentry (1978: 275) (as “Tynnanthus”). Type:—GUYANA. “Upper Mazaruni River Basin,
Kamarang River, Singh line from Akapai to Eboropu escarpment”, 13 October 1960, S.S. Tillett & C.L. Tillett 45643
(holotype MO! (2242100); isotypes K! (barcode 449550), NY! (barcode 328985), US! (barcode 125788), VEN!
(barcode 194023) photo).
Fig. 17: G–K
Lianas. Branchlets tetragonal to terete, without ritidome, finely striated, densely lenticeled, densely pubescent to
puberulent, with simple and peltate trichomes; interpetiolar ridge absent or present; interpetiolar patelliform glands
present; prophylls of the axillary buds 1.4–2.3 mm long, 1.1–2.3 mm wide, minute, shallowly triangular to
triangular, densely puberulent throughout, with simple and peltate trichomes. Leaves 2–3 foliolated (more
commonly 3); terminal leaflets often modified into trifid tendrils, without adhesive-disks on tip; petioles and
petiolules densely pubescent to puberulent throughout surface, with simple, peltate and patelliform trichomes;
petioles 1.5–7.5 cm long; petiolules 0.4–4 cm long; leaflets 5.9–15.4 cm long, 3.1–11.3 cm wide, chartaceous to
coriaceous, discolor, obovate to elliptic; apex acuminate or obtuse, mucronate; base cuneate or obtuse,
symmetrical; margin entire; the abaxial surface densely pubescent to puberulent (sometimes glabrescent)
throughout, with simple, peltate and patelliform trichomes; the adaxial surface pubescent to glabrescent,
throughout (sometimes only on and near the veins), with simple, peltate and patelliform trichomes; glandular
trichomes distributed especially on the abaxial surface; second venation weak brochidodromous or
brochidodromous; pocket domatia with or without trichomes. Inflorescence axilar or terminal, a thyrse or a
compound thyrse, lax, with conical aspect, first order (1.8–)3.6–6.3 cm long, second order 1.5–3.5 cm long; axis
densely pubescent or densely puberulent, with simple and peltate trichomes; bracts of the inflorescence
predominantly caducous, densely pubescent to pubescent throughout, 0.6–1.5 mm long; floral bracts 0.3–1 mm
long; floral pedicels 1–10 mm long. Calyx green to yellowish, 3–4.5 mm long, 2.5–4 mm wide, with transversal
aperture, minutely 5-denticulate (sometimes truncate), densely pubescent to puberulent throughout outside, without
patelliform glands; lobes 0.1–0.4 mm long. Corolla white, cream or pale yellow (sometimes pale lilac), 1–1.6 cm
long, 4–6 mm wide at the tube opening; tube 5–8 mm long, internally tomentose at the base, with simple and long
and short stipitate trichomes; nectar guides present, yellow; lobes densely pubescent to pubescent throughout lower
ones and at the margin of upper ones; upper ones 0.6–1.4 mm long, 1.2–2.4 mm wide, acute to obtuse; lower ones
3–5.1 mm long, 3–6 mm wide, obtuse to rounded. Androecium with fertile stamens inserted 2–3 mm from the base
of the corolla; shorter ones 6–9 mm long; longer ones 7–10 mm long; anthers thecae 0.9–1.2 mm long, obovate to
elliptic, subexserted; connective extending 0.1–0.3 mm beyond anther attachment; staminode 2–2.6 mm long,
glabrescent, with long and short stipitate trichomes. Gynoecium 11–12 mm long; ovary 1.5–1.7 mm long, 0.9–1.2
mm wide, conical, densely pubescent; style 8–11 mm long, tomentose to pubescent at the base. Fruit a linear
flattened capsule, 20–55 cm long, (1.2–)2.3–4.2 cm wide, woody, granular throughout (sometimes smooth near the
midvein), without lenticels to densely lenticeled, densely pubescent to pubescent, with simple, peltate and
patelliform trichomes; central ridge double, prominent to very prominent; margins prominently raised (winged),
0.5–1.4 cm wide. Seeds body 1.2–2.8 cm long, 0.9–1.3 cm wide; wings 1–1.8 cm long.
Phenology:—Flowers from August to December and produces fruits from February to July.
Distribution and habitat:—Occurs in moist broadleaf forests from Brazil (Acre, Amazonas, Pará and
Rondônia), French Guiana, Peru (San Martín), Suriname (Brokopondo and Nickerie) and Venezuela (Amazonas)
(Fig. 19).
Additional specimens examined:—BRAZIL. Acre: Sena Madureira, Anjo, 5 September 1997, I. Miranda et
al. 1657 (INPA). Amazonas: Presidente Figueiredo, Rebio Uatumã, grade do PPBio, 9 June 2007, J.G. Carvalho-
Sobrinho & J.R. Mesquita 1563 (INPA). Pará: Região do Rio Jari, Estrada entre Monte Dourado e Munguba, km
4, 15–16 May 1969, N.T. Silva 1966 (IAN, MO, NY); Ibid., Estrada do Munguba, km 7, 21 May 1969, N.T. Silva
2023 (MO, NY); Ibid., Estrada entre Planalto A e Tinguelim, Km 16, 11 July 1969, N.T. Silva 2398 (MO, NY).
Entre Acará e Moju, 2 September 1975, R.R. Santos 398 (IAN). Belém, Mocambo, Embrapa Forest Reserve, ca. 10
km from Belém, Transect 6, near sea level, 14 November 1984, A.H. Gentry 49049 (IAN, MO). Itaituba, Rio
Tapajós, S. Luiz, 4 December 1919, A. Ducke s.n. (R 22493, RB barcode 58493). Monte Dourado, Rio Jari, atrás do
aviário, 12 June 1968, E. Oliveira 4491 (IAN, NY). Oriximiná, Rio Trombetas, margem esquerda ao lado do lago
Muçura a 30 km de Porto Trombetas, 23 July 1980, C.A. Cid et al. 1748 (INPA, MG, NY). Santarém, Estrada para
o Rio Curuá Una, acampamento do Guaraná, 30 November 1966, P. Cavalcante & M. Silva 1474 (MG photo, MO,
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Phytotaxa 216 (1) © 2015 Magnolia Press
NY); Rio Curuá Una, acima da Cachoeira do Palhão, 8 December 1966, P. C a v a l c an t e & M . S i l v a 1 6 3 6 (IAN).
Rondônia: Eixo JP 14, 20 May 1987, S. Romaniuc Neto et al. 686 (SP). Porto Velho, Instituto Agronômico do
Norte, 1952, E. Cordeiro & J.F. da Silva 162 (MO, RB). FRENCH GUIANA: Station des Nouragues, Bassin de
l'Arataye, 7 July 1989, D. Sabatier & M.F. Prévost 2616 (P). Saül and vicinity, Route de Bélizon, N of Eaux
Claires, 200–400 m, 10 August 1993, S. Mori et al. 23213 (NY); Ibid., less than 500 meters S of Eaux Claires,
230–300 m, 19 September 1994, S. Mori et al. 23971 (NY, P, US). PERU. San Martín: Mariscal Cáceres, Tocache
Nuevo, Trail up Río Huallaga Valley toward Limón, 500 m, 11 March 1979, A.H. Gentry et al. 25566 (F photo,
MO); 11 March 1979, A.H. Gentry et al. 25579 (F, MO, NY). SURINAME. Brokopondo: NW Brokopondo
Stuwmeer Lake (E of Brownsberg Nature Reserve), Tonka Island, trail west from main compound, 15 m, 11
February 1998, M.J. Plotkin et al. 1293 (MO); Ibid., 15 m, 4 February 1999, B. Hoffman et al. 5282 (MO).
Nickerie: Area of Kabalebo Dam project, ca. 4 km S of road camp, ca. 23 km SW of Avanavero dam site, 18
November 1976, N.M. Heyde & J.C. Lindeman 164 (MO). VENEZUELA. Amazonas: Forest edge along Puerto
Ay ac uc ho –S am ari ap o hig hw ay 16 –2 5 km S o f Pu ert o Ayac uc ho , 10 0 m, 28 J un e 19 75 , A.H. Gentry & P. Berry
14503 (MO). Parcela de estudio del mamure al E del Raudal Gavilán, 100 m, 1 February 1991, G.A. Romero et al.
2289 (MO). Transecta entre Raudal Gavilancito, Río Gavilán, y la parcela de mamure, incluyendo zona al N del
caño que bodea la parcela, 80–100 m, 9 February 1992, G.A. Romero et al. 2331 (MO).
FIGURE 19. Distribution of Tyna nth us pub esc ens .
Taxo no mi c no te s: Tynanthus pubescens is morphologically similar to T. den sif loru s (see taxonomic notes
under T. densiflor us), and also shares obovate to elliptic leaflets with T. cogna tus (see taxonomic notes under T.
cognatus). Tynanthus pub escens is sister to another Amazonian species, T. panure nsis (Medeiros & Lohmann
2015), with which it also shares other morphological features (see taxonomic notes under T. panurensis). Despite
those morphological similarities, T. pubescens can be easily recognized by the densely pubescent to puberulent
branchlets, petioles and petiolules, interpetiolar patelliform glands, and fruits with double central ridge.
13. Tynanthus sastrei Gentry (1980: 214). Type:—FRENCH GUIANA. Sinnamary: “Route de Ste. Elie, 3 km avant
la parcelle, ARBOCEL”, 23 September 1977, C. Sastre 6015 (holotype MO! (2630185); isotypes CAY! photo, P!
(barcodes 481495, 481496)).
Fig. 20: A–D
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TAXONOMIC REVISION OF TYNANTHUS (BIGNONIEAE)
FIGURE 20. A–D. Ty na nt hu s sa st rei : A. Flowering branch; B. Interpetiolar region showing the foliaceous prophylls of the axillary
buds; C. Flower; D. Calyx with denticulate apex (L. Skog 7043, NY). E–J. T. sch uman nia nus : E. Flowering branch; F. Interpetiolar
region showing the bromeliad-like prophylls; G. Detail of inflorescence axis indumentum, showing patelliform trichomes; H. Open
corolla showing the androecium; I. Open calyx showing the gynoecium (M. Nee 38171, NY); J. Fruit (M. Lewis 37532, MO).
Illustration by Klei Sousa.