Phytotaxa 147 (1): 13–25 (2013) ISSN 1179-3155 (print edition)

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Copyright © 2013 Magnolia Press Article PHYTOTAXA
ISSN 1179-3163 (online edition)

http://dx.doi.org/10.11646/phytotaxa.147.1.2

Notes on the genus Swartzia (Leguminosae) in Ecuador, with descriptions of two

new species

BENJAMIN M. TORKE1* & ÁLVARO J. PÉREZ2

1
Institute of Systematic Botany, The New York Botanical Garden, Bronx, NY, 10458-5126, USA.
E-mail: btorke@nybg.org
2
Laboratorio de Ecología de Plantas, Herbario QCA, Escuela de Ciencias Biológicas, Pontificia Universidad Católica del Ecuador,
Apartado 17-01-2184, Quito, Ecuador. E-mail: ajperezc@puce.edu.ec
* Corresponding author

Abstract

Herbarium- and field-based research indicates that the neotropical genus Swartzia is represented in Ecuador by at least
17 species. A key is provided to aid their identification. In addition, two species, previously undocumented in the
literature, are described and illustrated. Both appear to be endemic to Ecuador. Swartzia decidua is known from a
confined area of the pre-montane humid forest zone on the western slopes of the Andes Mountains in Pichincha
Province, where it is threatened by habitat destruction. It belongs to S. sect. Paucistaminae and is notable in the context
of the genus for its very large petal and deciduous phenology. Swartzia yasuniensis occurs in Amazonian rainforest in
the drainage of the Napo River in Orellana Province and is a new member of S. sect. Pittierianae. In an intensively
studied 25-hectare plot in Yasuni National Park, it averaged 4 individuals (≥1 cm dbh) per hectare.

Resumen

Investigación basada en trabajo de campo y de herbario indica que el género neotropical Swartzia está representado en
Ecuador por lo menos por 17 especies. Se presenta una clave para ayudar con su identificación. Además, dos especies
previamente indocumentadas en la literatura, son descritas e ilustradas. Ambas parecen ser endémicas para Ecuador.
Swartzia decidua es conocida de un área confinada de la zona del bosque húmedo premontano en la vertiente occidental
de la Cordillera de los Andes en la provincia de Pichincha, donde se ve amenazada por la destrucción del hábitat.
Pertenece a S. sec. Paucistaminae y es notable en el contexto del género por su pétalo de gran tamaño y por ser árboles
caducifolios. Swartzia yasuniensis ocurre en selva amazónica en la cuenca del río Napo en la provincia de Orellana y es
un nuevo miembro de S. sec. Pittierianae. En una parcela de 25 hectáreas intensamente estudiada en el Parque Nacional
Yasuní, se promedió 4 individuos (≥ 1 cm dap) por hectárea.

Key words: endemic, Fabaceae, neotropic, Papilionoideae, taxonomy

Introduction

Swartzia (Leguminosae), a neotropical genus of some 200 species of trees and shrubs, is well represented in
the wet lowlands of Ecuador on both sides of the Andean Cordillera. Although there are many exceptions, the
genus is characterized by red-oxidizing exudate in the secondary phloem, distichous, imparipinnate leaves, a
stipellate leaf rachis, opposite lateral leaflets with brochidodromous secondary veins, monopetalous flowers, a
zygomorphic androecium, dimorphic, relatively numerous stamens, and arillate seeds (Cowan, 1968; Torke &
Mansano, 2009). It was weakly supported as monophyletic in the molecular phylogenetic analysis of Torke &
Schaal (2008).

Accepted by Vidal Mansano: 16 Oct. 2013; published: 20 Nov. 2013 13

 As part of an ongoing taxonomic study of the genus throughout its range, collections from Ecuador, both
in domestic herbaria and abroad, were examined. Seventeen species of Swartzia, including several
undescribed species, were found to be present in the country. While this estimate represents a modest increase
from the number previously documented (see: Cowan 1968, 1985, Neill & Palacios 1989, Neill et al. 1999,
Renner et al. 1990, Neill & Ulloa Ulloa 2011), purported records of several species —S. auriculata Poeppig
(1845: 61), S. benthamiana Miquel (1851: 15), S. cardiosperma Bentham (1870: 33), S. laevicarpa Amshoff
(1939: 37) S. leptopetala Bentham (1840: 87), and S. recurva Poeppig (1845: 61)—were found to be based on
misidentified specimens and/or outdated taxonomic concepts (see Torke 2007). Three species appear to be
endemic to the country, while another four are confined to the Pacific lowlands of northwestern Ecuador and
the adjacent Colombian Department of Nariño.
Here, formal descriptions are provided for two newly discovered endemic species, along with a key to the
species of Swartzia that occur in Ecuador. Three of the species included in the key remain undescribed and
will be dealt with in separate publications pending additional research. Conservation assessments using IUCN
Red List categories are based on the criteria and recommendations of the IUCN (2001) and the IUCN
Standards and Petitions Subcommittee (2011).

Systematic Treatment

Swartzia decidua Torke & A. J. Pérez, sp. nov. (Fig. 1)

Bracteolate pedicels, a large, cordate-based, yellow petal, three larger stamens, and an elongate gynoecium with an
oblong-linear ovary suggest a close phylogenetic relationship between the new species and the species of Swartzia
sect. Paucistaminae Torke & Mansano (2009: 921), to which it is assigned; in the context of the section, it is singular
in the following combination: bracts 1–1.5 mm long, pedicels 2.9–5.5 cm long, petal 6.6–8.4 cm wide, ovary 2.7–3.1
mm wide, densely sericeous-lanate, the locule densely lanate.
Type:—ECUADOR. Pichincha: [Mun. Pedro Vicente], Reserva Forestal ENDESA, Río Silanche: “Corporación Forestal
Juan Manuel Durini”, km 113 de la carretera Quito–Puerto Quito, faldas occidentales, a 10 km al N de la carretera
principal, 0°5’N, 79°2’W, 650–700 m elev., 7 April 1984 (fl), J. Jaramillo 6576 (holotype QCA!; isotypes AAU!,
NY!).

Deciduous tree, flowering prior to and during leaf initiation, to ca. 25 m tall; pubescence mostly ferruginous,
of simple trichomes, mostly 0.2–0.8 mm long, the shorter hairs dense, often somewhat twisting, tangled or
matted, the longer hairs less dense and relatively straight, these erect to sub-appressed; current season, leaf-
bearing branchlets 4.4–5.4 mm thick at middle of internodes, densely tomentose. Leaves imparipinnate, with
4–5 pairs of opposite lateral leaflets; stipules 7.3–11.3 × 5.2–8.5 mm, ovate, apically acute, glabrous adaxially,
densely tomentose abaxially, tightly clustered along new, unextended shoots; petioles 3.1–7.1 cm long, 2–2.8
mm thick at middle, terete to subterete, densely tomentose, the pulvinus ca. 6–9 × 3.1–4.3 mm; rachis 16–23
cm long, 1.1–2.6 mm thick at middle of segments, densely tomentose, longitudinally bicarinate along
segments adaxially, the ridges terminating in stipels, these ca. 0.6–2.1 mm long, triangular to ovate-lanceolate,
obscured by dense pubescence; petiolules 1.9–5.5 × 1.9–2.6 mm, densely tomentose; laminas 1.8–2.8 × longer
than wide, 5.5–16.5 × 2.8–8.4 cm, membranous and somewhat translucent in newly expanded leaves,
becoming chartaceous and opaque with maturity, the basal ones elliptic to ovate, the distal ones elliptic to
obovate or weakly oblong, the base usually obtuse or rounded, often broadly acute in the terminal leaflet, the
apex usually obtuse or rounded and then shortly acuminate, the adaxial surface for the most part glabrescent
but tomentose on the primary vein and margin, the abaxial surface fairly densely pilose, villous-tomentose on
the major veins, the venation immersed adaxially, prominent abaxially, the secondary veins 12–18 on each
side of the primary vein, initially ascending at 25°–35°, progressively upward-curving, fading but loosely
brochidodromous submarginally. Inflorescences simple racemes, apparently borne from annotinous or older
branches, to ca. 15-flowered; axes 11–18 cm long, 3.3–4.3 mm thick at base, densely tomentose; bracts ca. 1–

14 • Phytotaxa 147 (1) © 2013 Magnolia Press TORKE & PÉREZ

1.5 mm long and equally wide, broadly triangular, abaxially convex, glabrous adaxially, densely villous-
tomentose abaxially, rapidly caducous; pedicels 2.9–5.5 cm long, 2.7–2.8 mm thick at middle, somewhat
dorso-ventrally compressed, a trifle dilated toward apex; bracteoles 1.6–1.9 × 0.6–0.9 mm, narrowly
triangular-lanceolate, densely villous-tomentose; flower buds 10–11.7 × 6.2–10.6 mm, ellipsoid, shortly
umbonate, densely villous-tomentose. Calyx segments 3–5 in number, ca. 9.5–15 × 5.5–9.5 mm, deflexed,
glabrous adaxially, densely villous-tomentose abaxially. Corolla monopetalous; petal 5.8–6.5 × 6.6–8.4 cm,
yellow, with the venation somewhat darker, broadly reniform, essentially sessile, with the claw reduced to a
short, broad, obtusely triangular bulge between the lobes of the cordate base, this ca 0.5 × 7 mm, essentially
glabrous adaxially, but with a few hairs on the central part of base, fairly densely golden-sericeous to
tomentose on the central base and primary veins abaxially, more sparsely so between them, the venation
palmate with ca. 20 primary veins. Androecium zygomorphic, the stamens dimorphic, of two size classes;
larger stamens usually 3, occasionally 2, abaxial, the filaments ca. 19–24 mm long, 1.2–1.4 mm thick at base,
dorso-ventrally compressed, yellow, dilated and villous toward base, the anthers 2.4–2.9 × 1.1–1.4 mm,
oblong in outline, glabrous; smaller stamens ca. 140–160 in number, adaxial, glabrous, the filaments ca. 7–16
× 0.1–0.2 mm, terete to somewhat dorso-ventrally compressed, the anthers 0.7–1.3 × 0.9–1.3 mm, oblate to
elliptic in outline, the base and apex indented between the thecae. Gynoecium unicarpellate; ovary stipe 8.5–
12 × 1.4–1.6 mm, more or less terete, somewhat dilated at base and apex, densely whitish to tannish sericeous-
lanate; ovary proper 19–22 × 2.7–3.1 mm, arcuate, oblong-linear in outline, somewhat compressed laterally,
densely whitish to tannish sericeous-lanate, the locule densely lanate; ovules ca. 9; style 8.3–11.3 × ca. 0.7
mm, terminal, terete, linear, sericeous-lanate at base glabrous toward apex; stigma truncate. Mature fruits
unknown, but probably moniliform.
Distribution, habitats and conservation:—Swartzia decidua is known from only two collections from
Pichincha Province that were gathered between 650 and 700 m elevation in the upper Blanco-Guayllabamba
drainages, tributary systems of the Esmeraldas River, on the western slope of the Andean Cordillera (Fig. 2).
The localities from which the collections were made lie within a much larger zone that was historically
dominated by humid pre-montane forest, with associated tree species including Pouteria capacifolia Pilz,
Protium ecuadorense Benoist, Carapa spp., Brosimun utile (Kunth) Oken, Virola dixonii Little, Iriartea
deltoidea Ruiz & Pav., and Wettinia quinaria Burret (Jørgensen & Ulloa Ulloa, 1989).
The paucity of existing collections prevents concrete assessment of the conservation status of the species.
However, we suspect that its status is very tenuous given that: 1) the extent of occurrence (EOO, criterion B1)
is extremely small (problematic to estimate from only two collections, but perhaps <100 km2 and very likely
<5000 km2); 2) the species is known from only two closely positioned localities and has not been collected
since 1984; and 3) the extent of the sub-montane humid forest on the Pacific slope has been drastically
reduced by deforestation during the past few decades, and what little is left is highly fragmented and
increasingly degraded. Therefore, the species is provisionally assigned to the IUCN Redlist category of
Endangered (EN). Ongoing threats to the known habitat include dairy farming activities, palm heart (Bactris
gasipaes Kunth) cultivation, and extraction and commercialization of timber and non-timber forest products
(Altamirano, 2012). Zeas (2011) estimated that less than 1300 hectares of natural vegetation remains in Pedro
Vicente Municipality, where the two existing collections were gathered. One of the collections was made in a
privately protected, 85-hectare reserve owned and managed by ENDESA, an Ecuadorian forestry company
(see Jørgensen & Ulloa Ulloa, 1989); its conservation may well be critical for the preservation of the species.
Phenology:—Specimen label notes describe the species as being deciduous (Oldeman 3398), hence the
epithet. According to the collector, flowering commences during a leafless phase prior to the onset of new
leaves. Although the two specimens, both with anthesis flowers, were collected at different times of the year
—in April and November, respectively— on both the leaves appear to be immature.
Additional specimen examined (paratype):—Ecuador. Pichincha: [Mun. Pedro Vicente], vía Puerto
Quito, km 120, 660 m elev., 18 November 1975 (fl), R. A. A. Oldeman 3398 (QCA, U).

SWARTZIA (LEGUMINOSAE) IN ECUADOR, WITH TWO NEW SPECIES Phytotaxa 147 (1) © 2013 Magnolia Press • 15
FIGURE 1. Swartzia decidua. A: Branchlet and young leaves with detail of portion of adaxial leaflet surface showing trichomes and a
second detail of petiole bases and stipules; B: Defoliate branchlet with emerging shoots not yet elongated; C: Sub-mature
inflorescence; D: Inflorescence; E: Pedicel and flower bud (note subopposite bracteoles). A, D, E from Jaramillo 6576 (NY); B, C
from Oldeman 3398 (U); drawn by Bobbi Angell.

16 • Phytotaxa 147 (1) © 2013 Magnolia Press TORKE & PÉREZ

FIGURE 2. Geographical distributions of S. decidua and S. yasuniensis.

Discussion:—The new species is referred to the small section Paucistaminae, based on its possession of
bracteolate pedicels, a large, cordate-based, yellow petal, relatively few larger stamens and an elongate
gynoecium with the ovary more or less linear and substantially longer than the stipe and style. The section is
distributed from Panama and Colombia to northern Peru, where it occurs in humid lowlands on both sides of
the Andes Mountains, and also in the low-elevation valleys that penetrate the Andean Cordilleras. Swartzia
decidua differs from consectional species in its densely sericeous-lanate ovary with a densely pubescent
locule, smaller bracts, and longer pedicels (Table 1). It is nearly unique in the genus in being apparently
deciduous, the condition also occurring in the distantly related species S. cubensis (Britton & P. Wilson in:
Britton, 1926: 460) Standley (1935: 61) and S. pittieri Schery (1952: 263) (B. M. Torke, unpubl. data; N. L.
Cuello, pers. comm.), as well as in at least some populations of the closely related species S. macrosema
Harms (1926: 970) (van der Werff 19416). Deciduousness is usually associated with seasonal climates and is
an uncommon phenomenon in highly aseasonal habitats such as the humid pre-montane forest from which S.

TABLE 1. Selected diagnostic characters of the species of Swartzia sect. Paucistaminae.

No. Rachis
Bract Pedicel Bracteole Petal Ovary Ovary
lateral wings
Species length length length width width Ovary indument locule
leaflet presence/
(mm) (cm) (mm) (cm) (mm) indument
pairs absence
S. decidua 4–5 – 1–1.5 2.9–5.5 1.6–1.9 6.6–8.4 2.7–3.1 Densely Densely
Torke & A. J. Pérez sericeous-lanate lanate
S. haughtii (5–) 6–8 – (+) 1.5–4.9 0.8–2.3 0.9–3 2.9–4.1 1.2–1.4 Glabrous to thinly Glabrous
R. S. Cowan sericeous
S. macrosema Harms 3–6 – 5.4–12 0.9–3.1 1.1–6.8 4.2–7.9 1.4–2.6 Glabrous Glabrous
S. robiniifolia Willd. 6–10 + 3–7.2 0.7-1.8 3–7 2.7–3.5 1.4–1.8 Glabrous Glabrous
ex Vogel

SWARTZIA (LEGUMINOSAE) IN ECUADOR, WITH TWO NEW SPECIES Phytotaxa 147 (1) © 2013 Magnolia Press • 17
decidua was collected. Moreover, the observation that cycles of flowering followed by leaf flush occur at
disparate times of the year in S. decidua, as indicated by the existing collections, suggests that the
evolutionary origin of deciduousness in the species may have been driven by factors other than seasonality.
One hypothesis is that it is an adaptation to enhance visibility and/or accessibility of the inflorescence to
potential pollinators.

Swartzia yasuniensis Torke & A. J. Pérez, sp. nov. (Figs. 3, 4)

Ebracteolate pedicels, a nearly glabrous gynoecium, with the ovary ca. 3.5 × longer than wide, and a frequently branched
inflorescence belie a close phylogenetic relationship between the new species and several species of Swartzia
section Pittierianae (R. S. Cowan 1968: 24) Torke & Mansano (2009: 921), to which it is assigned; in the context of
the section, it is singular in the following combination: stipules 1.6–13 mm long, abaxial leaflet surface pilose,
bracts 3.1–6.6 mm long, buds 6.4–7.1 mm long, calyx segments 4–6, larger stamens abaxial, smaller stamens 74–95,
ovary stipe 4–6.5 mm long, ovary 6.1–8.1 mm long, ovules ca. 14.

Type: ECUADOR. Orellana: Estación Científica Yasuní, Rio Tiputini, al NO de la confluencia con el Río Tivacuno, 6
km E de la carretera Maxus, km 44, desvio hacia el pozo Tivacuno, Sendero Botánico, 0°38’S, 76°30’W, 200–300 m
elev., 7 November 1995 (fl), K. Romoleroux & R. Foster 1958 (holotype: QCNE-130955!; isotypes: F!, QCA!).

Evergreen tree to ca. 35 m tall and 56 cm dbh; pubescence mostly of golden or ferruginous, erect to sub-
appressed, fairly straight to weakly twisting, simple trichomes, these mostly 0.1–0.7 mm long; current season,
leaf-bearing branchlets 1.2–3.5 mm thick at middle of internodes, densely pilose-tomentose. Leaves
imparipinnate, with 3–5 pairs of opposite to subopposite lateral leaflets; stipules 1.6–13 × 0.8–2.3 mm,
narrowly lanceolate, widest at base, often falcate, glabrous adaxially, densely villous-tomentose abaxially,
glabrescent, caducous; petioles 0.8–3.2 cm long, 1.1–2.2 mm thick at middle, terete to subterete,
longitudinally bicarinate adaxially toward apex, densely pilose-tomentose, the pulvinus 2.7–5.8 × 1.4–2.5
mm; rachis 3.5–18 cm long, 0.6–2.2 mm thick at middle of segments, densely pilose-tomentose,
longitudinally bicarinate along segments adaxially, the ridges terminating in stipels, these 0.3–1.7 mm long,
triangular to subulate, densely sericeous; petiolules 1.3–3 × 1–2.1 mm, pilose; laminas (1.5–) 2–3.5 × longer
than wide, (1.9–) 4.2–14.4 × (0.9–) 1.9–5.5 (–7.6) cm, chartaceous, the basal ones smaller and less elongate
than the others, usually elliptic, the medial and distal ones elliptic, obovate or weakly oblong, the base acute,
obtuse or rounded, the apex usually acuminate, sometimes acute or obtuse, the tip rounded, the adaxial surface
mostly glabrous, pilose on the primary vein and margin, the abaxial surface often somewhat glaucous or
canescent, pilose, fairly densely so on the major veins, the primary vein impressed and other venation
immersed adaxially, all venation prominent abaxially, the secondary veins ca. 6–12 on each side of the
primary vein, initially ascending at 35°–55°, curving upward, fading or weakly brochidodromous
submarginally. Inflorescences racemes, frequently compound, with 1–2 orders of branching, borne from leaf
axils or from annotinous branchlets just below leaves, to ca. 50-flowered; axes (1.3–) 2.7–15.5 cm long, ca.
1.2–2.2 mm thick at base, densely pilose-tomentose; bracts 3.1–6.6 × 1.6–3.3 mm, elliptic to obovate,
abaxially convex, the apex acute or shortly acuminate, glabrous adaxially, densely sericeous-tomentose
abaxially, caducous; pedicels 4–12 mm long, 1.2–1.6 mm thick at middle, dorso-ventrally compressed,
densely pilose-tomentose; bracteoles absent; flower buds 6.4–9 × 4.5–7.1 mm, ellipsoid, shortly umbonate,
densely sericeous-tomentose. Calyx segments 4–6 in number, 2.4–5.7 mm wide, recurved, glabrous adaxially,
densely sericeous-tomentose abaxially. Corolla monopetalous, the petal yellow, mostly glabrous, sparsely
sericeous on the base of the primary veins abaxially; claw ca. 1.2 mm long, 0.4 mm wide at base, 1.1 mm wide
at apex; limb membranous, broadly elliptic, ca. 9.5 × 8.2 mm, the venation palmate, with ca. 7 primary veins.
Androecium zygomorphic, the stamens dimorphic, most of two size classes; larger stamens 4–5 in number,
abaxial, often with 1–2 smaller and less robust than the others, the filaments ca. 9–11.5 × 0.3–0.5 mm,
somewhat dorso-ventrally compressed, dilated basally, yellow, glabrous to sparingly villous, the anthers 1.2–
1.6 × 0.6–0.8 mm, elliptic or oblong-elliptic in outline, light tan, glabrous; smaller stamens ca. 74–95 in

18 • Phytotaxa 147 (1) © 2013 Magnolia Press TORKE & PÉREZ

number, adaxial, glabrous, the filaments ca. 4–7 × 0.1 mm, terete to somewhat dorso-ventrally compressed,
yellow, the anthers 0.8–1.1 × 0.6–0.8 mm, elliptic in outline, light tan. Gynoecium unicarpellate, dull yellow;
ovary stipe 4–6.5 mm long, 0.4–0.6 mm thick at middle, more or less terete, dilated at base and apex,
sparingly villous; ovary proper 6.1–8.1 × 1.8–2.3 mm, arcuate, inequilaterally and narrowly elliptic in outline,
laterally compressed, sparingly villous on the abaxial (dorsal) suture, otherwise glabrous, the locule glabrous;
ovules ca. 14; style 1.8–2.2 mm long, 0.3–0.4 mm thick at middle, terminal, terete, glabrous; stigma rounded-
truncate, green. Fruits 1–several-seeded, glabrous; stipe 9–17 × 1.7–2.8 mm, more or less terete, somewhat
dilated basally; body ca. 7–18 × 2.1–2.6 cm, inequilaterally elliptic to oblong-lanceolate in outline, laterally
compressed, the valves little or not at all constricted between seeds, the base and apex acute to somewhat
acuminate, the surface smooth to superficially rugose, with the ridges longitudinal to oblique.
Distribution, habitats and conservation:—Swartzia yasuniensis is known only from the northwestern
section of Yasuní National Park, thus the epithet, and the adjacent Huaorani Ethnic Reserve in the province of
Orellana in the Ecuadorian Amazon, where it occurs at less than 400 m elevation in the upper drainages of the
Tiputini and Yasuní Rivers, southern tributaries of the Napo River (Fig. 2). The actual distribution is probably
larger, as much of the surrounding region has been little collected. The species occurs in humid lowland
tropical rainforest, often in hilly dissected terrain, with a thin organic layer over red clay. According to
specimen label notes (Jaramillo & Tapia 18679A; Pérez & Santillán 4682), associated tree species include
Astrocaryum chambira Burret, Brownea grandiceps Jacq., Cedrelinga catenaeformis (Ducke) Ducke, Iriartea
deltoidea Ruiz & Pav., Iryanthera hostmannii (Benth.) Warb., Parkia nitida Miq., Protium nodulosum Swart,
and Rinorea lindeniana (Tul.) Kuntze (see also: Valencia et al. 2004).
The new species is one of several species of Swartzia found in the well-studied 50-hectare Yasuní Forest
Dynamics Plot. In 25-hectares of the plot, there were 103 individuals of Swartzia yasuniensis with dbh greater
than or equal to 1 cm, an average of four individuals per hectare. Of these, eight individuals exceeded 30 cm
dbh, while 21 individuals had a dbh exceeding 10 cm but less than 30 cm. Between 2002 and 2007, the annual
mortality rate for the species in the plot was 0.8%; recruitment rate was 0.4 individuals per year, and average
growth rate was 0.9 mm per year.
Swartzia yasuniensis is assigned to the IUCN Redlist category of Near Threatened (NT). The documented
extent of occurrence (EOO, criterion B1) is quite small (less than 5000 km2), and on its periphery, encroaching
settlement and deforestation are apparent in satellite images. However, the larger region is still mostly covered
by intact and poorly explored Amazonian plant communities, and the species is present in at least two large,
nationally protected areas.
Phenology:—Flowering has been observed in November, fruiting in January.
Additional specimens examined (paratypes):—ECUADOR. Orellana: Parque Nacional Yasuní,
carretera y oleoducto de MAXUS, km 40, parcela permanente #10 de 1 hectárea, árbol marcado #06.21,
0°39’S, 76°26’W, 250 m elev., 15–17 May 1994 (st), M. Aulestia 2223 (MO, NY, QCNE); Carretera
Pompeya, Sur-Iro–Parque Nacional Yasuní, km 38.7, junto a la Estación de Monitoreo de Fauna Onkonegare
Ecuambiente S. A., 0°39’31.7”S, 76°27’7.5”W, 200–300 m elev., 26 June 1994 (st), X. Buitrón et al. 632
(QCA); Cant. La Joya de Los Sachas, Yasuní National Park, km 14–15 of the Maxus pipeline road, E of Juan
Tapuy's finca, Plot Shipati 3, tree #458, 0°31’S, 76°32’W, 250 m elev., October 1997 (st), N. Pitman & T.
Delinks 2448 (QCA); Parque Nacional Yasuní, km 9 via NPF–Pozo Tivacuno, 0°38’S, 76°30’W, 220 m elev.,
7 June 1997 (st), J. Jaramillo & I. Tapia 18679A (QCA); Estación Científica Yasuní, Parcela de 1 ha, km 8.2
de la carretera NPF–Tivacuno, 0°40’51”S, 76°23’12”W, 200–300 m elev., 11 February 2002 (st), J. Jaramillo
et al. 23061 (QCA); Estación Científica Yasuní, parcela de 1 ha, km 8.2 de la carretera NPF–Tivacuno,
0°40’51”S, 76°23’12”W, 200–300 m elev., 1 March 2002 (st), J. Jaramillo et al. 23257 (QCA); Parque
Nacional Yasuní, km 34.6, carretera Pompeya-Iro, parcela 6, 0°37.637’S, 76°27.736’W, 250–300 m elev., 12
August 1997 (st), M. J. Macía et al. 1135 (QCNE); Parque Nacional Yasuní ECY, Sendero “Laguna,”
0°40’40”S, 76°23’40”W, 200–300 m elev., 28 January 2010 (fr), A. J. Pérez & W. Santillán 4682 (QCA);
Añangu, Parque Nacional Yasuní, 0°31’–0°32’S, 76°23’W, 260–350 m elev., 30 May–21 June 1982 (st), SEF
8869 (QCA, QCNE, US); 21 June 1982 (st), SEF 8993 (QCA, QCNE).

SWARTZIA (LEGUMINOSAE) IN ECUADOR, WITH TWO NEW SPECIES Phytotaxa 147 (1) © 2013 Magnolia Press • 19
FIGURE 3. Swartzia yasuniensis. A: Leafy branchlet and inflorescence, with detail of stipule; B: Apex of inflorescence with
submature flower buds subtended by bracts; C: Leafy branchlet with inflorescence; D: anthesis flower, lateral view; E: Petal, adaxial
view; F: Post-anthesis flower with fertilized gynoecium; G: Submature fruit, fresh; H: Submature fruit, dried; I: Leaf. A, B, E, F from
Romoleroux 1958 (F); C, D, G, from photo by Á. J. Pérez; H. from Pérez & Santillán 4682 (QCA); I from Macía et al 1135 (QCNE);
drawn by Bobbi Angell.

20 • Phytotaxa 147 (1) © 2013 Magnolia Press TORKE & PÉREZ

FIGURE 4. Swartzia yasuniensis. A: Leafy branchlet with inflorescence; B: Leaf, abaxial view; C: Flowers; D: Infructescences; E:
Sub-mature fruits; photographed by Álvaro J. Pérez.

Discussion:—Swartzia yasuniensis is assigned to the small section Pittierianae (R. S. Cowan 1968: 24)
Torke & Mansano (2009: 921), which comprises five species. Sectional characters include the lack of
bracteoles, a glabrous or nearly glabrous gynoecium, a relatively compact ovary and a frequently branched
inflorescence. Among consectional species, it differs from S. pittieri Schery (1952: 263) and S. trianae
Bentham (1870: 39) of the northern Andes, the Orinoco basin, and northwestern Amazonia most obviously in
having more numerous calyx lobes, larger floral bracts, and typically more densely pubescent leaflets (Table
2). Probably more closely related are S. jorori Harms (1915: 39) and S. juruana Torke (2004: 358) of
southwestern Amazonia, the former also ranging southward to the Brazilian Pantanal, but it differs from them
in its longer stipules, floral bracts and ovary stipe, more strongly zygomorphic androecium, more numerous
smaller stamens and ovules, and more frequently multi-seeded fruits (Table 2).
The documented distribution of S. yasuniensis is similar to that of S. bombycina R. S. Cowan (1985: 303).
Both species appear to be restricted to the so-called Napo ecoregion in the Ecuadorian Amazon, giving
credence to the classification of the ecoregion as a distinct biogeographical province (e.g., Silva et al. 2005).

SWARTZIA (LEGUMINOSAE) IN ECUADOR, WITH TWO NEW SPECIES Phytotaxa 147 (1) © 2013 Magnolia Press • 21
TABLE 2. Selected diagnostic characters of the species of Swartzia sect. Pittierianae.

Abaxial No. Larger Ovary Ovary

Stipule Bract Bud No.
leaflet calyx stamen stipe proper No.
Species length length length smaller
indu- seg- orien- length length ovules
(mm) (mm) (mm) stamens
ment ments tation (mm) (mm)
S. jorori Harms 0.6–3 Nearly 0.8–2.5 3.4–5.8 2–4 Radial 14–39 0.8–2.5 3.3–5.5 3–7
glabrous
S. juruana Torke 1.1–2.5 Pilose to 2.2–3.1 5.7–8.8 3–4 Radial 36–59 1.9–2.2 5.8–7 5–8
strigose
S. pittieri Schery 2.8–12.1 Nearly 1–1.7 3.7–5.2 2 Abaxial 50–65 3.8–5.9 5.5–8 11–13
glabrous
S. trianae Benth. 1.6–8.2 Glabrous 1.3–3.6 8–11.3 2 Abaxial 108–125 5.7–11 12.6–25 14–17
to pilose
S. yasuniensis 1.6–13 Pilose 3.1–6.6 6.4–9 4–6 Abaxial 74–95 4–6.5 6.1–8.1 ca. 14
Torke & A. J. Pérez

A key to the species of Swartzia occurring in Ecuador

1. Petal white, violet or pink; leaves multifoliolate; mature fruits often more than 3 cm broad; Amazonian Region ..... 2
- Petal yellow (or the flowers apetalous); leaves multifoliolate or unifoliolate; mature fruits less than 3 cm broad;
Amazonian region, Andean slopes, and Pacific lowlands ............................................................................................ 5

2. Ovary and ovary stipe densely pubescent, the fruits usually retaining at least some pubescence on the stipe, sutures
and/or valves ................................................................................................................................................................ 3
- Ovary and stipe glabrous or essentially so, the fruits entirely glabrous ...................................................................... 4

3. Stipules 2–32 mm long; abaxial surface of leaflets drying with secondary and higher-order veins darker than or sim-
ilar in color to the lamina; calyx appreciably sericeous-strigose on the adaxial surface, particularly toward center ....
............................................................................................................................................ S. bombycina R. S. Cowan
- Stipules not apparent or not exceeding 1 mm long; abaxial surface of leaflets usually drying with the secondary and
higher order veins discernibly lighter than the lamina; calyx essentially glabrous on the adaxial surface ...................
................................................................................................................................................ S. rosea Mart. ex Benth.

4. Leaflet venation raised-reticulate on both surfaces; inflorescence axes, pedicels and flower buds glabrous; flower
buds 10.4–19.4 mm long .............................................................................................................. S. calva R. S. Cowan
- Leaflet venation immersed and inconspicuous on both surfaces; inflorescence axes, pedicels, and flower buds strigu-
lose; flower buds 4.2–8.1 mm long ....................................................................................................S. polyphylla DC.

5. Leaves unifoliolate....................................................................................................................................................... 6
- Leaves multifoliolate ................................................................................................................................................. 10

6. Pedicels supplied with a pair of bracteoles (not to be confused with stipules that sometimes subtend bracts); ovary
stipe at least twice as long as the ovary proper; fruits maturing green, leaflets strongly discolorous; trunk slash with
red-oxidizing exudate.......................................................................................................S. klugii (R. S. Cowan) Torke
- Pedicels lacking bracteoles; ovary stipe not more than 1.5 × the length of the ovary proper; fruits maturing orange,
yellow, green or brownish-green; trunk slash lacking red-oxidizing exudate ............................................................. 7

7. Androecium semi-actinomophic, the stamens more or less isomorphic, or at least not clearly divided into two size
classes; flower buds 3.2–5.4 mm broad; petal 4.4–12.3 mm broad, the base of the limb acute to truncate...................
........................................................................................................................................ S. arborescens (Aubl.) Pittier
- Androecium strongly zygomorphic, the stamens dimorphic, with an abaxial cluster of larger stamens and an adaxial
or central cluster of smaller stamens; flower buds 5.8–12.2 mm broad; petal 16.1–45.9 mm broad, the base of the
limb cordate ................................................................................................................................................................. 8

22 • Phytotaxa 147 (1) © 2013 Magnolia Press TORKE & PÉREZ

8. Leaf stalk almost entirely pulvinular, auriculate on the adaxial side just below apex, but not obviously winged; leaf-
let blade often more than 2.5 × longer than wide; inflorescences frequently borne from defoliate nodes of branches,
but also from leaf axils; widespread in Amazonian lowlands .................................................... S. calophylla Poeppig
- Leaf stalk usually with a short “rachis” between the pulvinus and pulvinulus, this typically marginate or winged;
leaflet blade usually less than 2.5 × longer than wide; inflorescences borne mostly in the axils of current leaves;
Pacific lowlands, frontal ranges, and Andean slopes ................................................................................................... 9

9. Larger stamens 8–15; leaflets acute to obtuse at base, the secondary veins raised to plane on the adaxial surface;
eastern Andean slope, Cordillera del Condor, and other eastern slope frontal ranges ............ S. simplex (Sw.) Spreng.
- Larger stamens 17–25; leaflets often rounded or subcordate at base, the secondary veins usually noticeable
impressed on the adaxial surface; Pacific lowlands and western Andean slope ...............S. aff. simplex (Sw.) Spreng.

10. Pedicel supplied with a pair of bracteoles (not to be confused with stipules that sometimes subtend bracts); larger
stamens 2–3 ................................................................................................................................................................ 11
- Pedicel lacking bracteoles; larger stamens 2–ca. 18 or the stamens essentially isomorphic..................................... 13

11. Lateral leaflets mostly 6–8-paired; petal 2.9–4.1 cm wide ..................................................... S. haughtii R. S. Cowan
- Lateral leaflets mostly 3–6-paired; petal 4.2–8.4 cm wide ....................................................................................... 12

12. Ovary densely sericeous-lanate; bracts 1–1.5 mm long; pedicels 2.9–5.5 cm long; western Andean slope .................
......................................................................................................................................S. decidua Torke & A. J. Pérez
- Ovary glabrous; bracts 5.4–12 mm long; pedicels 0.9–3.1 cm long; eastern Andean slope and adjacent Amazonia....
..................................................................................................................................................... S. macrosema Harms

13. Primary vein raised-cariniform on the adaxial leaflet surface; lateral leaflets 1–3-paired; leaf rachis winged; flower
buds and abaxial surface of calyx glabrous ................................................................................................................ 14
- Primary-vein impressed or immersed on the adaxial leaflet surface; lateral leaflets 2–6-paired; leaf rachis usually
unwinged; flower buds and abaxial surface of calyx pubescent ................................................................................ 15

14. Androecium semi-actinomophic, the stamens more or less isomorphic, or at least not clearly divided into two size
classes; flower buds 3.2–5.4 mm broad; petal 4.4–12.3 mm broad; gynoecium unicarpellate; fruit maturing green or
brown-green; Amazonian lowlands ............................................................................... S. arborescens (Aubl.) Pittier
- Androecium strongly zygomorphic, the stamens dimorphic, with an abaxial group of ca. 9–18 larger stamens and an
adaxial or central group of much more numerous smaller stamens; flower buds 7.5–12 mm broad; petal 25–53 mm
broad; gynoecium often composed of two to several free carpels; fruit maturing orange or yellow; Pacific lowlands
and Cordillera del Condor … S. littlei R. S. Cowan

15. Ovary excluding the stipe densely pubescent; fruits retaining at least some pubescence on the stipe, valves or sutures
................................................................................................................................................................................... 16
- Ovary excluding the stipe glabrous or nearly so; fruit glabrous ................................................................................ 17

16. Lateral leaflets 2–3-paired; flower buds 7–11 mm broad; ovary 12–18 mm long; Pacific lowlands ............................
..................................................................................................................................................S. aff. amplifolia Harms
- Lateral leaflets 4–6-paired; flower buds 4.4–5.2 mm broad; ovary 6–7.5 mm long; Amazonian lowlands .................
................................................................................................................................................ S. aff. leptopetala Benth.

17. Calyx segments 2; ovary 12.6–25 mm long; petal 4–5 cm broad .......................................................S. trianae Benth.
- Calyx segments 4–6; ovary 6.1–8.1 mm long; petal less than 1 cm broad ....................................................................
.................................................................................................................................S. yasuniensis Torke & A. J. Pérez

Acknowledgments

The authors thank Bobbi Angell for preparing the illustrations, Wayne Law, manager of the GIS Lab at the
New York Botanical Garden, for preparing the map, Renato Valencia and Consuelo Hernandez for providing
data from the 50-ha Yasuni Forest Dynamics Plot, and the curators of the herbaria cited for making their

SWARTZIA (LEGUMINOSAE) IN ECUADOR, WITH TWO NEW SPECIES Phytotaxa 147 (1) © 2013 Magnolia Press • 23
specimens available for study. Funding was provided by the National Science Foundation (DEB-0918498),
the BIOYAS Project, and the “Impuesto a la Renta” taxes of the Pontificia Universidad Católica del Ecuador
(PUCE). Permission for AJP’s work in Yasuni National Park was granted by the Ecuadorian Ministerio del
Ambiente.

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