Ladenbergia siranensis (Rubiaceae: Cinchoneae), a new species

from the Sira Mountains, Eastern Andes of Central Peru, and the identity
of Ladenbergia acutifolia
EDER A. CHILQUILLO TORRES1,2, ANDRÉ OLMOS SIMÕES3, AND JOAQUINA ALBÁN CASTILLO2
1
Programa de Pós-Graduação em Biologia Vegetal, Instituto de Biologia, Universidade Estadual de
Campinas, Av. Monteiro Lobato 255, Campinas, SP CEP: 13083-970, Brazil; e-mail:
e.chilquillo.torres@gmail.com
2
Departamento de Etnobotánica, Museo de Historia Natural, Universidad Nacional Mayor de San Marcos,
Av. Arenales 1256, Jesús María, Lima, Peru; e-mail: jalbanc@gmail.com
3
Departamento de Biologia Vegetal, Instituto de Biologia, Universidade Estadual de Campinas, Av.
Monteiro Lobato 255, Campinas, SP CEP: 13083-970, Brazil; e-mail: andreosimoes@gmail.com

Abstract. A new species of Ladenbergia (Rubiaceae) has been found in the Cordillera de Sira
in the eastern Andes of central Peru. It is here described as Ladenbergia siranensis, and it is
morphologically similar to L. acutifolia, L. graciliflora and L. shawistigma. It differs from
these by a shrubby habit, coriaceous, narrowly elliptic leaves, few-flowered inflorescences, the
hypanthium hirtellous outside, small and deeply lobed calyx, a glabrous style, and fruits with
reddened hirtellous pubescence. A key to these four species is provided.
Keywords: Neotropics, outlying range, sclerophyllous vegetation, taxonomy.

The Cordillera El Sira is an isolated mountain
range situated between the Pachitea and Ucayali
rivers in the easternmost outlying range of the
Andes mountains, in Central Peru, Huánuco Re-
gion (Terborgh & Weske, 1975). This mountain
range reaches a maximum elevation of ca. 2500
m, exhibiting 7000 mm of annual rainfall (Kuijt &
Graham, 2015), and has been described as a
unique ecosystem, because of its potential for
yielding new taxa and its relatively high number
of endemic species of vertebrates (Terborgh &
Weske, 1975; Duellman & Toft, 1979; Harvey
et al., 2011) and plants (Monteagudo et al., 2014).
Early botanical explorations in the Cordillera
El Sira were conducted by Theodore Dudley in
1969, as part of a larger program to measure the
distribution of avian species along the Altitudinal
Transect Yuyapichis (ATY) (Terborgh & Dudley,
1973). Later, an Austrian-Peruvian Expedition
directed by W. Morawetz in 1987–1988 revisited
the ATY and made plant collections that included
new species and new distributional records (e.g.,
Prance, 1995; Sastre, 1996; Grant, 2004;
Wasshausen, 2007). Recently, forest inventories
by RAINFOR (http://www.rainfor.org) in
permanent plots in the ATY have contributed
substantially to increasing the knowledge of its
tree flora (Monteagudo et al., 2014).
The neotropical genus Ladenbergia Klotzsch
belongs to tribe Cinchoneae of Rubiaceae
(Andersson, 1995a; Andersson & Antonelli,
2005), and comprises ca. 35 species (Andersson,
1997; Taylor & Gereau, 2010; Chilquillo, 2016),
distributed from Costa Rica to Bolivia. Its distri-
bution is centered in the Tropical Andes of which
the Peruvian Andes holds the highest diversity
with 14 species (Andersson, 1995b; Chilquillo
et al., 2017). Ladenbergia is distinguished from
other Rubiaceae genera by the following combi-
nation of features: tree or shrubby habit, terminal
inflorescences, heterostylous flowers, distally
ridged corolla lobes, capsules that are septicidal
with basipetal or acropetal dehiscence and numer-
ous flattened seeds with winged margins
(Andersson, 1994, 1997).
Ladenbergia is morphologically similar to
Cinchona L. and is usually confused with it in
the field. The two genera are similar in vegetative
characters and in inflorescence form and fruit, but
they differ in floral characters: Cinchona has pale

Brittonia, 71(2): 166Y171 (2019), DOI 10.1007/s12228-018-9562-0

ISSN: 0007-196X (print) ISSN: 1938-436X (electronic)
© 2019, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.

Published Online: 16 January 2019

2019] CHILQUILLO ET AL.: LADENBERGIA SIRANENSIS (RUBIACEAE)
to deep pink or purple-red corollas that are dense-
ly pubescent in the throat, while Ladenbergia has
white corollas or flushed with pink on the tube
base that are glabrous or shortly puberulous in the
throat (Andersson, 1995a).
During recent botanical exploration at the ATY,
the first author found several species of
Ladenbergia that were also collected by the
Austrian-Peruvian Expedition: L. klugii
Andersson, L. muzonensis (Goudot) Standl.,
L. graciliflora K. Schum., L. macrocarpa (Vahl)
Klotzsch), and an undescribed species that was
previously misidentified as L. acutifolia (Ruiz &
Pavon.) Klotzsch. Here, we describe and illustrate
the new species and compare it with similar spe-
cies of Ladenbergia.
Material and methods
This study is based on standard taxonomic
methods, and is part of a comprehensive revision-
ary study of Ladenbergia by the first author. The
following herbaria were consulted: CEN, CR, F,
G, GB, HOXA, INB, MO, MOL, NY, R, RB, UB,
UEC, USJ, USM. The digital databases and
photos of specimens from COL, CUZ, K, MA,
P, QCA, QCNE, US were also consulted. Termi-
nology was adapted from Andersson (1998). The
conservation status assessment was based on the
IUCN Red List categories and criteria (IUCN,
2012) and subsequent guidelines (IUCN
Standards And Petitions Subcommittee, 2017).
Taxonomic treatment
Ladenbergia siranensis Chilq., sp. nov. Type:
Peru, Huánuco: Cordillera del Sira, Provincia
Pachitea, elfin forest after the BCampamento
Peligroso^, 74°43’W, 9°25’S, 1700 m. elev., 1
May 1988, B. Wallnöfer 17-1588 (holotype:
GB; isotypes: W–unmounted, LZ–not seen).
(Fig. 1, 3B)
Diagnosis: Ladenbergia siranensis is distinguished from
other Ladenbergia species by its shrubby habit; narrowly el-
liptic, coriaceous leaves, few-flowered inflorescence; hypan-
thium hirtellous outside; small (4–4.7 mm long) and deeply
lobed calyx, corolla tube 12–15 mm long; glabrous style, and
fruits with reddened hirtellous indument.
Shrubs, 3–4 m tall. Young branches glabrous or
sparsely puberulous, with trichomes ferrugineous.
Stipules shortly fused at the base, oblong in the
167
basal portion with the upper part acuminate, gla-
brous to puberulous, 1–1.1 × 0.3–0.35 cm. Petioles
1.1–1.65 cm long, glabrous to sparsely puberulent.
Leaves opposite, blades plane, thick in texture
when fresh, chartaceous when dry, 6.5–10.2 ×
1.4–2.25 cm, narrowly elliptic (length/width ratio
4.1–4.6), acute at base, acute to acuminate at apex,
adaxial surface matte to subnitid and glabrous,
abaxial surface with midrib +/- hirtellous from
bristly hairs, intervenous surface glabrous or
sparsely and minutely strigulose, margins weakly
revolute, tuft domatia absent, secondary veins 6–8
pairs, weakly brochidodromous, slightly sulcate
above, prominent beneath, tertiary venation weak-
ly evident above and below with alternate-
percurrent arrangement. Inflorescence terminal,
branched to 1–2 orders, 5–9-flowered, broadly
obtriangular in outline, axes villosulous, peduncles
1–1.2 cm, rachis 0.8–1 cm, two-noded, with axes
dichasial, bracts linear-lanceolate, generally decid-
uous. Flowers 6-merous, nocturnal, strongly fra-
grant, sessile to subsessile with pedicels 0.5–
1.5 mm long, bracteoles 1–2.5 mm long.
Hypanthium obconic, pilosulous to tomentulose,
3–3.8 × 2.5–3 mm. Calyx 4–4.7 mm long, deeply
lobed, lobes acute, 3–3.6 mm long, puberulous to
hirtellous outside, glabrous to sparsely hirtellous
inside. Colleters not seen. Corolla salverform,
fleshy, white, tube 12–15 mm long, hirtellous out-
side, inside villous in upper half and glabrous in
lower half, lobes 12.5–14 mm long, ridged part
glabrous to puberulous, central part hirtellous.
Filaments 0.5–1.1 mm long and attached ca. 9–
10.5 mm above base of corolla tube (ca. 75% of
tube length). Anthers 3.9–4.1 mm. Style ca. 9 mm
long, glabrous, stigmatic lobes ca. 3.5 mm long.
Capsules woody, oblong-cylindrical, externally
puberulous to hirtellous, indument red/reddened,
30–35 × 4.5–5.5 mm, endocarp 0.4–0.5 mm thick.
Seeds flattened, oblong-fusiform, 11.5 × 4.1 mm,
with wings marginally fimbriate (Fig. 1).
Distribution and habitat.—Ladenbergia
siranensis is only known from summit of the Cor-
dillera El Sira, Pachitea, Huánuco, Peru (Fig. 2).
These mountains lack traces of human activities.
Their summit (1700–2000 m) is covered by
sclerophyllous vegetation, usually called Bbosque
enano^ (Vásquez et al., 2010), characterized by
the absence of trees and predominance of shrubby
plants (Ilex sp., Clethra castaneifolia Meisn., Styrax
vilcabambae (D. R. Simpson) B. Walln, Symplocos
quitensis Brand, Clusia sp., Gordonia fruticosa
(Schrad.) H. Keng, Gaultheria sp., Myciaria sp.,
168 BRITTONIA [VOL 71

FIG 1. Line drawing of Ladenbergia siranensis Chilq. A. Flowering branch. B. Stipules. C. Detail of venation leaf blade. D.
Flower and bud. E. Detail of the inside of the calyx. F. Detail of the inside of the corolla. G. Style. H. Fruit. I. Seed. (Drawn by Klei
Souza from B. Wallnöfer 17-1588, GB-6123490.)

Freziera sp.) (Monteagudo et al., 2014), growing on
sandstone with nonwoody vegetation (mosses, bro-
meliads, orchids, and sedges) (Kuijt & Graham,
2015) (Fig. 3A).
Etymology.—Ladenbergia siranensis is named
for the type locality, to call attention to the bio-
logical diversity and high endemism in the Cor-
dillera El Sira, and thereby, to encourage the
preservation of that biodiversity.
Phenology.—Collected with flowers in May
and fruits in September.
Conservation status.—Ladenbergia siranensis
is only known from four collections from only
one site in a large and poorly explored area situ-
ated within the Reserva Comunal El Sira (created
by Peruvian government DS. N 037-2001-AG).
Its location within this communal reserve affords
some protection, as does the steep topography and
2019] CHILQUILLO ET AL.: LADENBERGIA SIRANENSIS (RUBIACEAE) 169

FIG 2. Geographic distribution of Ladenbergia siranensis and morphologically similar species.

current lack of roads in the Sira which limit the
montane forest’s exposure to human influence.
Nevertheless, mining, logging, and oil explora-
tion are active in the region and threaten the
montane forests (Dourojeanni, 2015). As a con-
sequence, it is not possible to determine its con-
servation status, and we here provide a prelimi-
nary classification of data deficient (DD; see
IUCN, 2012; IUCN Standards and Petitions
Subcommittee, 2017).
Additional specimens examined. PERU. Huánuco: Cor-
dillera El Sira, Provincia Pachitea, Region Pucallpa,
elfin forest after the Campamento Peligroso, 13
Apr 1988, B. Wallnöfer 16-13488 (GB, LZ, W), 4 Sep
2017, E. Chilquillo et al. 5489 (USM), 5 Sep 2017,
E. Chilquillo et al. 5492 (USM).
Ladenbergia siranensis presents a shrubby
habit and is unique in the genus by the combina-
tion of the following characters: oblong stipules
that are shortly fused at the base; narrowly elliptic
(length/width ratio 4.1–4.6), thick-textured leaf
blades; inflorescences few-flowered with short
axes and peduncles; sessile to subsessile flowers,
calyx limb deeply lobed, corolla tube 12–15 mm
long, villous inside in the upper half, glabrous
styles; and capsules oblong-cylindrical with red
hirtellous indument (Fig. 3B).
Ladenbergia siranensis is morphologically
similar and usually confused with the poorly
known Peruvian species L. acutifolia. This latter
species is known only from two collections [Ruiz
& Pavon s.n (G, BM, MA, photo F-136); Schunke

FIG 3. A. Sclerophyllous vegetation on the summit of Cordillera El Sira. B. Infrutescence of Ladenbergia siranensis, frontal
view. (Photos: A, by Julio Monzón; B, by Eder Chilquillo.)
170 BRITTONIA [VOL 71

5310 (F, G, IAN, NY)]. It was treated differently
in the taxonomic synopsis of Ladenbergia by
Andersson (1997) than here. Recent collections
and field observations in eastern Peru found that
the additional collections cited as L. acutifolia by
Andersson (1997) actually correspond to the more
common species L. graciliflora [Belshaw 3464
(K, NY); Gentry et al. 37910 (MO)]. True
Ladenbergia acutifolia can be recognized by its
tree habit; ovate to oblong leaf blades
(length/width ratio 2.3–2.7) with cordate or
subcordate to rounded bases; multiflowered lax
inflorescence with two or three orders of
branching; and corolla tube 8–10 mm long.
Ladenbergia siranensis also resembles
L. shawistigma, which shares the shrubby habit,
coriaceous and elliptic to lanceolate leaf blades,
and deeply lobed calyx limb. However,
L. shawistigma differs from L. siranensis by its
unbranched umbelliform inflorescence.

Key to Ladenbergia siranensis, L. acutifolia, L. graciliflora and L. shawistigma

1. Inflorescence short and unbranched, in an umbelliform cyme with three flowers; style hirtellous. ……………L. shawistigma
1. Inflorescence short to well developed and lax, in dichasial cymes with five to many flowers; style glabrous.
2. Shrubs; leaves narrowly elliptic; inflorescence branched to one or two orders, with 5–9 flowers; fruits with hirtellous red
pubescence……….......……………………………………………………………………………………… L. siranensis
2. Treelets or trees; leaves elliptic to oblong; inflorescence branched to 2–4 orders, with 12–90 flowers; fruits glabrous or with
colorless puberulous pubescence.
3. Corolla tube 30–35 mm long; capsule narrowly oblong, 40–50 mm long ....…………………………….. L. graciliflora
3. Corolla tube 8–10 mm long; capsule pyriform or oblong, 15–30 mm long ………………………………... L. acutifolia

Acknowledgments Andersson, L. 1998. A revision of the genus Cinchona

We thank Charlotte M. Taylor and two anony-
mous reviewers for valuable comments that
helped improve the final version of the manu-
script; the curators of all herbaria cited in for
access to specimens, Claes Persson, Bruno
Wallnöfer, Joao A. M. Carmo, André V. Scatigna,
Robin Fernandez Hilario and Elluz Huáman for
their collaboration in the preparation of the man-
uscript; Armin Niesner, Julio Monzón, Carlos
Vasquez BMoro^, Clever Llagas and Estación
Biológica Panguana for assistance and collabora-
tion in the field; and Klei Souza for the excellent
illustration. The first author thanks CAPES for a
Ph.D. scholarship.
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