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Phytotaxa 289 (3): 247–255
http://www.mapress.com/j/pt/
Copyright © 2016 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Cássio van den Berg: 6 Dec. 2016; published: 29 Dec. 2016
http://dx.doi.org/10.11646/phytotaxa.289.3.4
247
Maxillaria tenebrosa, a new species in the Maxillaria variabilis group (Orchidaceae:
Maxillariinae) from Southwestern Ecuador
BOSCO JAVIER ZAMBRANO ROMERO1 & RODOLFO SOLANO-GOMEZ2
1Centro de Tenencia Orquiecuador y Gloxinias, Sucre y García Moreno, esquina frente al Parque central, 071050 Piñas, El Oro, Ecua-
dor; e-mail: bosco_escorpion@hotmail.com
2Instituto Politécnico Nacional, Centro Interdisciplinario de Investigación para el Desarrollo Integral Regional Unidad Oaxaca, Hor-
nos 1003, Santa Cruz Xoxocotlán, 71230, Oaxaca, México.
Abstract
Maxillaria tenebrosa, a new species discovered in Southwestern Ecuador is described and illustrated. Information concern-
ing its distribution, habitat, floral biology, and conservation status is provided. The new taxon is a member of M. variabilis
group or M. section Erectae, and it is compared with morphologically similar species such as M. acervata, M. caespitifica,
M. caucae, M. costaricensis, M. dichaeoides, M. ponerantha, M. procurrens, M. variabilis, and M. xanthorhoda.
Resumen
Se describe e ilustra Maxillaria tenebrosa una nueva especie descubierta en el suroccidente de Ecuador. Se proporciona
información sobre su distribución, hábitat, biología floral y estatus de conservación. El nuevo taxón es integrante del grupo
de M. variabilis o M. sección Erectae y es comparado con especies morfológicamente similares como M. acervata, M. caes-
pitifica, M. caucae, M. costaricensis, M. dichaeoides, M. ponerantha, M. procurrens, M. variabilis y M. xanthorhoda.
Key words: Maxillaria variabilis group, Maxillaria sect. Erectae, Maxillariella, Maxillariinae, semi-deciduous montane
forest, semi-deciduous premontane forest
Introduction
Maxillaria Ruiz & Pavón (1794: 116) is a Neotropical genus containing approximately 660 species (Govaerts et al.
2014). Members of Maxillaria typically grow as epiphytes, or rarely as terrestrials in cloudy, wet or dry forests, from
Mexico to Peru and Bolivia, including the Antilles (Dodson 2002). Maxillaria, in its broadened delimitation (Chase et
al. 2015, Schuiteman & Chase 2015), is distinguished from other related genera of the Maxillariinae Bentham (1881:
288) because the species present a combination of the following characteristics: leaves with conduplicate development,
one-flowered inflorescences that emerges from the base of the pseudobulbs, floral bracts longer than the length of the
pedicelate ovary, lip articulated at the base of the column, sclerotic fibers in flowers, and fruits with lateral dehiscence
(Atwood & Mora de Retana 1999, Whitten et al. 2007, Chase et al. 2015, Schuiteman & Chase 2015).
The phylogenetic relationships of the Maxillariinae, from DNA sequences data, have been analyzed by several
studies in order to define the generic circumscriptions, especially in Maxillaria, the most diverse group in the subtribe
(Ojeda et al. 2003, Singer & Koehler 2003, Sitko et al. 2006, Dathe & Dietrich 2006, Whitten et al. 2007, Blanco et al.
2007, Szlachetko et al. 2012). These studies have shown that Maxillaria, as previously considered, was a polyphyletic
group within a highly supported clade, which also includes groups such as Anthosiphon Schlechter (1920: 182),
Chrysocycnis Linden & Reichenbach (1854: 280), Cryptocentrum Bentham (1881: 325), Cyrtidiorchis Rauschert
(1982: 560), Mormolyca Fenzl (1850: 253), Pityphyllum Fenzl (1850: 253), and Trigonidium Lindley (1837: t. 1923)
(Chase et al. 2015). Whitten et al. (2007) recognized several monophyletic groups within this clade, later Blanco et
al. (2007) proposed them as genera within Maxillariinae and, on the other hand, they considered Maxillaria with
a narrow circumscription. However, Chase et al. (2015) and Schuiteman & Chase (2015) extended the Maxillaria
ZAMBRANO & SOLANO
248 • Phytotaxa 289 (3) © 2016 Magnolia Press
delimitation to include all the groups of the well-supported clade previously recognized by Whitten et al. (2007).
Additionally, the generic groups recently segregated from Maxillaria by Blanco et al. (2007) and Szlachetko et al.
(2012) were considered as sections of this genus. This way, Maxillaria becomes the most diverse and morphologically
heterogeneous group within Maxillariinae.
Maxillaria includes the group of M. variabilis Bateman ex Lindley (1837: t. 1986) and similar species, which
originally was named by Pfitzer (1889: 187) as Maxillaria sect. Erectae and characterized by its erect rhizomes and
laterally compressed pseudobulbs. In the phylogenetic analysis of Whitten et al. (2007), representatives of this group
formed a monophyletic clade for which the name Maxillaria variabilis group was proposed; then Blanco et al. (2007)
elevated this clade to generic level with the name of Maxillariella Blanco & Carnevali (2007: 527) and it was also
recognized by Szlachetko et al. (2012). However, Schuiteman & Chase (2015) included this group within its expanded
concept of Maxillaria with its original name, Maxillaria sect. Erectae. Even though with this delimitation Maxillaria
is now a very large and diverse genus, it is highly supported as a monophyletic group. So, in order to avoid the
proliferation of generic names, the delimitation that includes all them under a single name defined by cladistics criteria
is followed here. This way, Maxillariella is here considered a member of Maxillaria as Maxillaria sect. Erectae.
Species in this section are recognizable by the combination of some of the following traits: rhizome elongated between
adjacent pseudobulbs; pseudobulbs ancipitous provided with one or two leaves; leaf slightly leathery, linear, and
unevenly 2-lobed at the apex; the solitary single inflorescence that emerges from the developing pseudobulb; flowers
bell-like; floral bract much shorter than the ovary; sepals and petals recurved at their apices; lip entire or slightly
3-lobed, with a glossy callus; column arched with a well-developed foot; anther minutely papillose, and fruits with
lateral dehiscence (Atwood & Mora de Retana 1999, Blanco et al. 2007, Whitten et al. 2007, Whitten & Blanco 2011,
Szlachetko et al. 2012, Schuiteman & Chase 2015).
In Ecuador around 200 species of Maxillaria are known, which represent almost 30% of the total diversity of
this genus; in addition, 50 of these taxa are considered endemic in the country and about 20 species are members of
the section Erectae (Jørgensen & León-Yánez 1999, Dodson 2002, 2004, Ulloa & Neill 2005, Blanco et al. 2007,
Schuiteman & Chase 2015). Nonetheless, with the exception of Dodson (2002), the taxonomic studies for Maxillaria
are scarce in the country and therefore its richness could be increased. Recently, some specimens of Maxillaria flowered
in cultivation (Orquiecuador & Gloxinias collection). The morphological traits of these plants revealed that they belong
to M. sect. Erectae, showing morphological similarities with Maxillaria acervata Reichenbach (1855: 217), Maxillaria
caespitifica Reichenbach (1877: 73), Maxillaria caucae Garay (1967: 258), Maxillaria costaricensis Schlechter (1923:
232), Maxillaria dichaeoides Bennett & Christenson (2009: 26), Maxillaria ponerantha Reichenbach (1854: 17),
Maxillaria procurrens Lindley (1845: 383), Maxillaria variabilis and Maxillaria xanthorhoda Schlechter (1918: 279).
However, when the material cultivated was compared with these species, it was found that they corresponded to a
different and unknown taxon, which is here described and illustrated for the first time.
Materials and methods
The description of the new taxon was based on cultivated specimens at the orchid collection of Orquiecuador &
Gloxinias (Piñas, El Oro, Ecuador). A complete specimen was drawn using a stereoscopic microscope USB 50-500x
and a Nikon® D7100 camera. An individual plant, a flower and the dissected floral parts were photographed and
illustrated using a Nikon® D7100 camera. A digital lamina showing vegetative and floral morphology was done from
these photos, which were edited with Adobe Photoshop® CS4 and Jasc® Paint Shop Pro 8. Illustrations and digital
composition included a typical plant habit, views of the flower, inflorescence, flower dissection, details of the petal and
lip, pedicel, ovary, column, lip, anther cap, and pollinarium. The voucher specimens were deposited at QCNE herbaria
and particular collection of Orquiecuador & Gloxinias. Distribution map showing the known localities for the new
taxon was made using the software ArcGIS 10 (ESRI 2012).
Taxonomic treatment
Maxillaria tenebrosa Zambrano & Solano sp. nov. Figs. 1 and 2
A NEW MAXILLARIA FROM ECUADOR Phytotaxa 289 (3) © 2016 Magnolia Press • 249
FIGURE 1. Maxillaria tenebrosa Zambrano & Solano. A. Habit. B. Flower, lateral view. C. flower dissection. D. Ovary, lip and column,
lateral view. E. Clinandrium margin, ventral view. F. Anther. G. Pollinarium. Drawn by B.J. Zambrano based on the holotype.
ZAMBRANO & SOLANO
250 • Phytotaxa 289 (3) © 2016 Magnolia Press
Similar to Maxillaria variabilis, from which is distinguished by their pendent habit, inflorescences produced from the mature pseudobulb,
porrect petals, and entire lip.
Type:—ECUADOR. El Oro: Cantón Piñas, filo el Trigal, 1200 m, 8 May 2007, Zambrano B. 101 (Holotype in QCNE; isotype in Herb.
J. Zambrano [in spirit]).
Plant epiphytic, pendent, rhizomatous, 25–30 cm tall. Rhizome pendulous, cylindrical, 2–3 cm long between adjacent
pseudobulbs, covered by conduplicate, distichous, overlapping, chartaceous, persistent, acute sheaths. Roots flexuous,
slender, whitish, 1 mm diameter. Pseudobulbs ellipsoid, distichous, ancipitous, light green, smooth when young,
longitudinally striated when mature, 3.0–4.5 cm long, 1.8–2.0 cm wide, apically 1-foliate (rarely 2-foliate), subtended
by 3–6 foliaceous distichous, conduplicate, persistent, chartaceous, and overlapping sheaths, sometimes purple spotted.
Leaf sessile, elliptic to lanceolate-ensiform, obtuse, slightly coriaceous, arching, conduplicate at the base, obliquely
bilobed at the apex, light green, sometimes purple spotted on abaxial surface in young leaves, slightly blurred in the
mature ones, 16–20 × 1.2–2.0 cm. Inflorescence from the mature pseudobulb, 1–3 per pseudobulb; peduncle erect,
cylindrical, 15 mm long, 1.5 mm diameter, with 3–4 overlapping bracts similar to the floral ones, 8–10 × 2.0–2.5 mm
(5 mm when extended). Floral bracts conduplicate, acute, carinate, scarious, 6–7 mm long. Inflorescence one-flowered,
bell-like, more or less open, producing a slightly fetid odor; the sepals, petals, and lip dark purple, the column light purple
and lustrous on ventral surface, the anther and pollinia dark purple. Ovary pedicellate, slightly arching, cylindrical,
longitudinal sulcate, 20 mm long, 2 mm diameter. Dorsal sepal fleshy, oblong-oblanceolate, obtuse, slightly concave,
shortly apiculate, slightly recurved at apex, 5–7-nerved, 13.0 × 4.5–5.0 mm; lateral sepal fleshy, obliquely oblong-
lanceolate, acute, shortly apiculate, slightly reflexed at the apices, slightly concave at the base, 5–7-nerved, 14.0 × 5.0
mm. Petals fleshy, porrect, obliquely oblanceolate, acute, shortly apiculate, slightly recurved at apex, slightly concave,
erose along the margins, 5-nerved, 12.0 × 3.5 mm. Lip entire, fleshy, arching, conduplicate, articulate at the base of the
column foot, the blade oval-elliptic when extended, sub-rounded, retuse, pilose-papillose at apex, 7–8-nerved, abruptly
attenuate at base, 13.0 × 7.5 mm; the lateral margins erose, erect and undulate up to the middle third; with a callus on
the lower middle of the lip, prominent, thickened, elevated, subquadrate, lustrous. Column slender, arching, semiterete,
slightly 2-winged at the apex, 8.5–9.0 mm long, 2.5–3.0 mm wide at the stigma level, with a downward foot, ca. 3
mm long, light purple and dark purple spotted; clinandrium marginally ciliolate. Anther apical, ovoid, galeate, pilose-
papillose, ca. 2.3 × 2.3 mm. Pollinarium formed by 4 pollinia, in two subequal pairs, longitudinal ovoid, 2.0 × 1.5 mm,
attached to a semilunar viscidium with the arms downwards. Capsule narrowly ellipsoid, with a persistent perianth, 2.5
cm long (3 cm including the remnant column), ca. 5 mm diameter.
Distribution and habitat:—Up to now Maxillaria tenebrosa is only known from two localities from El Oro
province, southwestern Ecuador. This species grow between 600 and 1,300 m in elevation, in semi-deciduous
premontane and semi-deciduous montane forests (Fig. 3). The plant grows as an epiphyte on tree branches of Ficus sp.
and Mauria sp.
Floral biology:—In culture, M. tenebrosa has flowered from March to December. The dark purple flowers,
fringed sepals and petals, landing platform formed by the lip, short hairs at apex of lip and column, and fetid odor
emitted by the flowers suggests that this species has a sapromyophylous pollination syndrome (van der Pijl 1966,
Argue 2012). Flower scent is more intense from 10:00 and 15:00; then it decreases and again it is intensified in the
morning the next day. This pattern in the production of the floral aroma might regulate pollinator visitation, which
could be attracted looking for feeding or egg-deposition sites in the flower.
Etymology:—The specific epithet is from the Latin tenebrosus, “dark”, in reference to the flower color.
Additional specimens:—ECUADOR. El Oro: Cantón Piñas, parroquia Capiro, near sector Antoniopamba, 856
m, 2 June 2014, Zambrano B. 1390 (Herb. J. Zambrano, cultivated in Orquiecuador & Gloxinias); parroquia Capiro,
on the way up to Loma del Guayacan, 840 m, 2 June 2014, Zambrano B. 1408 (Herb. J. Zambrano, cultivated in
Orquiecuador & Gloxinias); parroquia Capiro, sector Conchicola, 650 m, 21 September 2015, Zambrano. B. 1698
(Herb. J. Zambrano, cultivated in Orquiecuador & Gloxinias).
Comments:—Maxillaria tenebrosa is a member of the M. variabilis group or Maxillaria sect. Erectae, where the
most similar species are: M. acervata, M. caespitifica, M. caucae, M. costaricensis, M. dichaeoides, M. ponerantha, M.
procurrens, M. variabilis, and M. xanthorhoda. The new taxon is most similar with the Mesoamerican M. variabilis,
but it is different by its procumbent habit (vs. pendent), inflorescences produced from the developing pseudobulb (vs.
from the mature one), flowers yellow or yellow with red stain (vs. dark purple), petals strongly recurved (vs. porrect),
and lip lightly 3-lobed (vs. entire) (Bateman 1837, Jiménez et al. 1998). On the other hand, M. variabilis has not been
reported for Colombia and Peru (Bennett & Christenson 1993, 1995, 1998, 2001, Dodson 2004, Trujillo 2014), so
the geographic disjunction among the southernmost populations of this species and that from M. tenebrosa allows
recognizing them as different taxa.
A NEW MAXILLARIA FROM ECUADOR Phytotaxa 289 (3) © 2016 Magnolia Press • 251
FIGURE 2. Maxillaria tenebrosa Zambrano & Solano. A. Habit. B. Apex. C. Inflorescences. D. Flower, lateral view. E. Flower dissection.
F. Detail of the petal (500x). G. Detail of the lip (500x). H. Pedicel, ovary, column and lip, lateral view. I. Anther cap and pollinarium. J.
Column apex, pollinarium and anther cap, lateral view. K. Detail of the clinandrium margin, lateral and ventral view (500x). Plate by B.J.
Zambrano based on cultivated specimens.
ZAMBRANO & SOLANO
252 • Phytotaxa 289 (3) © 2016 Magnolia Press
TABLE 1. Summary of principal differences between Maxillaria tenebrosa, M. acervata, M. caespitifica, M. caucae, M. costaricensis, M. dichaeoides, M. ponerantha, M. procurrens, M. variabilis
and M. xanthorhoda.
Characteristic Maxillaria
acervata
Maxillaria
caespitifica
Maxillaria
caucae
Maxillaria
costaricensis
Maxillaria
dichaeoides
Maxillaria
ponerantha
Maxillaria
procurrens
Maxillaria
tenebrosa
Maxillaria
variabilis
Maxillaria
xanthorhoda
Growth habit Erect Pendent Erect Procumbent Erect Erect Procumbent Pendent Procumbent Erect
Pseudobulb form Ovoid Fusiform to
cylindric
Ovoid Fusiform Ellipsoid Ellipsoid Ellipsoid Ellipsoid Fusiform to
ellipsoid
Oblong-
obovate
Rhizome segments
(long)
≥ 2 cm ≤ 1.5 cm ≤ 1.5 cm ≥ 2 cm ≥ 2 cm ≥ 2 cm ≥ 2 cm ≥ 2 cm ≥ 2 cm ≥ 2 cm
Leaf form Oblong-
lanceolate
Lanceolate Oblong-
elliptic
Lanceolate Oblong-
elliptic
Lanceolate to
elliptic
Elliptic Elliptic to
lanceolate-
ensiform
Elliptic to
lanceolate
Obtuse-
elliptic
Leaf apex Obtuse to
unequally 2-
lobate
Unequally 2-
lobate
Emarginate Obtuse to
unequally
2-lobate
Unequally
2-lobate
Acute to
unequally
2-lobate
Emarginate-
mucronate
Obliquely 2-
lobate
Obtuse to
unequally 2-
lobate
Acute
Inflorescence (Origin) Developing
pseudobulb
Developing
pseudobulb
Developing
pseudobulb
Developing
pseudobulb
Developing
pseudobulb
Developing
pseudobulb
Mature
pseudobulb
Mature
pseudobulb
Developing
pseudobulb
Mature
pseudobulb
Petal apex Porrect to
slightly
recurved
Strongly
recurved
Slightly
recurved
Recurved Porrect Recurved Strongly
recurved
Porrect Strongly
recurved
Strongly
recurved
Petal (Nervation) 5 5 3 5 3 3-4 5 5 5 5
Petal margin Entire Entire Entire Entire Entire Entire Entire Erose Entire Entire
Ovary length
(including the pedicel)
≥ 15 mm ≤ 10 mm ≤ 10 mm ≥ 15 mm ≥ 15 mm ≤ 10 mm ≥ 15 mm ≥ 15 mm ≥ 15 mm ≥ 15 mm
Lip form Entire to
slightly 3-
lobate
Entire to
slightly 3-lobate
Entire 3-lobate Entire 3-lobate Entire Entire Slightly 3-
lobate
Slightly 3-
lobate
Lip apex Obtuse or
rounded
Retuse Retuse Retuse to
emarginate
Emarginate Emarginate Rounded Retuse Retuse Emarginate
Lip margin Entire Entire Erose Entire Erose Entire to
erose
Erose Erose Entire Erose
Lip (long) ≤ 10 mm < 8 mm < 8 mm ≥ 10 mm < 8 mm < 8 mm ≥ 10 mm ≥ 10 mm ≥ 10 mm ≥ 10 mm
A NEW MAXILLARIA FROM ECUADOR Phytotaxa 289 (3) © 2016 Magnolia Press • 253
FIGURE 3. Known localities for Maxillaria tenebrosa in Southwestern Ecuador. Map by B.J. Zambrano.
The South Central American M. caucae, originally described as M. parvula (Schlechter 1924: 176), can be
distinguished by its erect habit (vs. pendent), ovoid pseudobulbs (vs. ellipsoid), oblong-elliptic leaves (vs. elliptic to
lanceolate-ensiform), petals slightly recurved at the apex and 3-veined (vs. porrect, 5-veined), and ovary ≤ 10 mm long
including the pedicel (vs. ≥ 15 mm) (Garay 1967). Among other similar Maxillaria, the Central American and South
American M. caespitifica is different by its caespitose habit (vs. rhizomatic), lanceolate leaves (vs. elliptic to lanceolate-
ensiform), yellow or greenish yellow flowers (vs. dark purple), sepals 5–7 mm long (vs. 13–14 mm), strongly recurved
petals (vs. porrect), and lip < 8 mm long (vs. ≥ 10 mm). The south Central American and South American M. ponerantha
differs by its erect habit (vs. pendent), shorter leaves (2.5–8 cm vs. 16–20 cm), yellow-cream with red stain flowers (vs.
dark purple), petals ≤ 7 mm, 3–4-veined (vs. 12 mm, 5-veined). Another similar species is M. costaricensis, restricted
to Costa Rica and Panama, it differs from M. tenebrosa by its fusiform to cylindric pseudobulbs (vs. ellipsoid), green
to cream or rose flowers (vs. dark purple), and slightly 3-lobed lip (vs. entire) (Atwood & Mora de Retana 1999). The
South Central American M. acervata different by its shorter leaves (1.5–6 cm vs. 16–20 cm), green to white flowers,
red stained lip (vs. dark purple), marginally entire petals (vs. erose), and lip ≤ 10 mm, marginally entire and rounded
ZAMBRANO & SOLANO
254 • Phytotaxa 289 (3) © 2016 Magnolia Press
at apex, (vs. ≥ 10 mm, erose, and retuse) (Reichenbach 1855). The North Andean M. dichaeoides differs by having
oblong-elliptic leaves (vs. elliptic to lanceolate-ensiform), smaller, marginally entire and 3-veined petals (7 mm vs. 12
mm, marginally erose and 5-veined), lip < 8 mm long, without a callus (vs. ≥ 10 mm, with a lustrous callus) (Bennett &
Christenson 2009). The South American M. procurrens is distinguished from M. tenebrosa by its procumbent habit (vs.
pendent), leaves with apex mucronate (vs. non-mucronate), marginally entire petals (vs. erose), lip with a rounded apex
(vs. retuse) (Lindley 1845). Finally, M. xanthorhoda, endemic to Peru, differs by its rhizome of 5–6 cm long between
adjacent pseudobulbs (vs. 2–3 cm long), elliptic leaves (vs. elliptic to lanceolate-ensiform), pink with red spots flowers
(vs. dark purple), strongly recurved petals (vs. porrect), and lip with an emarginated apex (vs. retuse) (Schlechter 1918,
Bennett & Christenson 1998). Table 1 present a summary of the differences among the species mentioned above.
Conservation status:—Not evaluated, but it is desirable to consider this species in a risk category to ensure
its protection. Maxillaria tenebrosa has a very restricted geographical distribution. The only known localities are in
remnant forest disturbed by human activities, where the populations are at risk as a result of deforestation and their
low densities (1–3 plants per phorophyte). Furthermore, due to the fragmentation process that affects the habitat of the
known populations (Filo de Trigal and Capiro), they are isolated, little interconnected among them, out of protected
areas and so far, the species has not been found in other similar forests of the region.
Acknowledgments
An anonymous reviewer for its helpful and constructive comments, to the Ecuadorian Ministry of Environment (MAE)
by the permission authorized for this research. The present work is part of the project Diversidad y distribución
geográfica de la familia Orchidaceae del cantón Piñas, provincia de El Oro (N.◦006-IC-FLO-DPAEO-MAE).
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