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Synopsis of Ecuadorian Pterichis
(Orchidaceae)
Marta Kolanowska
1,2
, Dariusz L. Szlachetko
3
and Sławomir Nowak
3
1Department of Geobotany and Plant Ecology, Faculty of Biology and Environmental Protection,
University of Lodz, Lodz, Poland
2Department of Biodiversity Research, Global Change Research Institute AS CR, Brno,
Czech Republic
3Department of Plant Taxonomy and Nature Conservation, Faculty of Biology, University of
Gdansk, Gda
nsk, Poland
ABSTRACT
A taxonomic synopsis of the orchid genus Pterichis in Ecuador is presented.
All national representatives of this genus are characterized and their floral segments
are illustrated. Four new species and two new varieties are described. An updated key
to Ecuadorian Pterichis is provided. Plants of the genus are growing mostly as
terrestrial herbs at the altitude of 2,300–4,110 m. Often two or more species co-occur
in the area of 25 × 25 km. Their occurrence was reported from three ecoregions—the
Eastern Cordillera real montane forests, the Northern Andean páramo and the
Northwestern Andean montane forests. Seven Ecuadorian Pterichis are endemic.
Subjects Biodiversity, Biogeography, Plant Science, Taxonomy
Keywords Biodiversity, Cranichideae, Neotropics, New species, Taxonomy, Distribution
INTRODUCTION
Amazonia and the Andes are some of the richest diversity hotspots in the world
(Pennington et al., 2010;Antonelli et al., 2018). An extraordinary number of rare and
endemic species is observed especially in the area where the two regions meet—the
Tropical Andes. This hotspot extends from western Venezuela to northern Argentina and
Chile, and incorporates significant part of territories of Colombia, Ecuador, Peru, and
Bolivia. At the same time this is one of the most endangered ecoregions on the planet
(Mittermeier et al., 1999), with a large portion of its landscape having been transformed.
The fertile inter-Andean valleys of Colombia and Ecuador, are the most degraded as a
result of agriculture and urbanization (Zador, 2015). The further habitat loss constitute a
serious threat to numerous orchid genera (Hirtz, 2011).
Recognizing species composition is essential to preserve wildlife by identifying areas
important for biodiversity and ecosystem services (Sofo, 2011;Casetta, Marques da Silva &
Vecchi, 2019). For that reason, taxonomists are continuously publishing articles not only
introducing the novelties in local biota, but also more comprehensive articles presenting
concepts of particular plant or animal group (Ormerod, 2018;Damián Parizaca & Torres
Montenegro, 2018). Orchidaceae and Asteraceae, with a rate of 500 new species being
described each year (Chase et al., 2015), are among the most problematic groups to
work with.
How to cite this article Kolanowska M, Szlachetko DL, Nowak S. 2021. Synopsis of Ecuadorian Pterichis (Orchidaceae). PeerJ 9:e10807
DOI 10.7717/peerj.10807
Submitted 27 August 2020
Accepted 30 December 2020
Published 9 February 2021
Corresponding author
Marta Kolanowska,
martakolanowska@wp.pl
Academic editor
Mike Thiv
Additional Information and
Declarations can be found on
page 41
DOI 10.7717/peerj.10807
Copyright
2021 Kolanowska et al.
Distributed under
Creative Commons CC-BY 4.0
While globally about 2/3 of Orchidaceae are epiphytic or lithophytic plants, according
to the International Union for Conservation of Nature almost half of the extinct orchids
are terrestrial herbs (Swarts & Dixon, 2009). Unfortunately, no assessment of threats to
tropical terrestrial/lithophytic orchids was done so far. The evaluation of the diversity of
these plants and providing tools useful for their identification is essential to recognize the
most valuable and biodiverse areas and to undertake any conservation actions.
Here we are dealing with the diversity of the mostly terrestrial genus Pterichis in
Ecuador—one of the most orchid-rich countries in the Tropical Andes (Dodson, 2003;
Meisel & Woodward, 2005). Despite the relatively long history of studies on the
Ecuadorian Orchidaceae flora (Garay, 1978;Dodson & Dodson, 1980–1989;Dodson &
Luer, 2005;Dodson & Luer, 2010), novelties are reported from this country each year
(Zambrano Romero & Solano, 2019;Hágsater & Santiago, 2019).
As currently recognized, Pterichis encompasses ca. 45 species distributed from
Costa Rica and Jamaica to Bolivia with the greatest diversity observed in Colombia
(Kolanowska & Szlachetko, 2017). This genus was proposed in 1840 by Lindley to
accommodate plants characterized by a mix of features of Cranichideae (helmet-shaped
lip) and Spirantheae (plant habit). Representatives of Pterichis are usually terrestrial herbs
with tuberous, clustered roots. Their scape is loosely sheathed and leaves are rosulate
and often absent during flowering. The flowers are non-resupinate and arranged in a
loosely to densely flowered raceme. Their sessile, concave, fleshy lip is usually densely
papillate or/and ornamented with a series of swollen, knob-like cells along the margins
(Kolanowska & Szlachetko, 2017).
Garay (1978) reported the occurrence of six Pterichis species from Ecuador considering
P. seleniglossa as synonym of P. triloba,P. barbifrons as conspecific with P. parvifolia and
P. costaricensis as synonym of P. habenarioides. In the last few years four additional species
were described from this country (Szlachetko & Kolanowska, 2014;Kolanowska &
Szlachetko, 2015;Kolanowska & Szlachetko, 2019).
The generic separateness of Pterichis has never been argued. The genus is clearly defined
by its morphological characters—i.a. non-resupinate flowers with sessile, very broad lip
which is concave in the center, variously ornamented with papillae or knob-like
appendages. The monophyletic character of Pterichis was confirmed in the molecular
studies (Salazar et al., 2003,2009) and taxonomists agree with its placement in
Cranichideae (Dressler, 1981;Dressler, 1993;Szlachetko, 1995;Chase et al., 2015).
The aim of this study was to evaluate the diversity of Pterichis species occurring in
Ecuador, to provide key for their identification and to discuss the most discriminative
morphological characters of each species.
MATERIALS AND METHODS
Pterichis specimens deposited in Ecuadorian herbaria HA, LOJA, Q, QAP, QCA, QCNE,
and QPLS were examined in addition to material from AAU, AMES, COL, CUVC, F,
FLAS, FMB, G, HUA, K, LPB, MO, NY, P, PSO, RPSC, UGDA, VALLE, and W. Specimens
collected outside Ecuador were used as reference material to evaluate diagnostic usefulness
of particular morphological features. Specimens were examined during visits in almost all
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 2/45
of these facilities. Only material from AAU was loaned. Herbaria acronyms used in this
article follow Thiers (2020). A total of over 100 Ecuadorian collections of Pterichis were
examined.
The form of the leaf (if present) as well as the length and the surface of the scape were
studied in the beginning. The assessment of the vegetative structures included also the
tubular sheaths on the scape and the elements of the inflorescence (e.g., form of the floral
bracts and ovaries). The perianth segments were observed after softening flowers in boiling
water using a stereoscopic microscope. A database of original diagnoses and illustrations
published by authors of particular taxa was created.
Morphological features presented in this study were compiled based on Ecuadorian
material. The information provided on herbarium specimens labels was used for
georeferencing. Distribution maps were prepared using ArcGIS 9.3 (Esri, Redlands, CA,
USA). Species richness was calculated using DIVA-GIS (Hijmans et al., 2001). The concept
of ecoregions follows Olson et al. (2001).
Nomenclature
The electronic version of this article in portable document format will represent a
published work according to the International Code of Nomenclature for algae, fungi, and
plants (Turland et al., 2018), and hence the new names contained in the electronic version
are effectively published under that Code from the electronic edition alone. In addition,
new names contained in this work that have been issued with identifiers by IPNI will
eventually be made available to the Global Names Index. The IPNI Life Science
Identifiers (LSIDs) can be resolved and the associated information viewed through any
standard web browser by appending the LSID contained in this publication to the prefix
“http://ipni.org/.”The online version of this work is archived and available from the
following digital repositories: PeerJ, PubMed Central, and CLOCKSS.
RESULTS
We confirmed 17 Pterichis species in Ecuador. Seven taxa, including four new species
described here, are endemic—P. ansaloniana, P. dodsoniana, P. elliptica, P. hirtziana,
P. madsenii, P. meirax, and P. tunguraguona.
They are growing almost always as terrestrial herbs on roadsides, in cloud forest,
humid montane forest and paramo at the altitudes of 2,300–4,110 m (Fig. 1).
The broadest altitudinal range is observed in P. acuminata (2,300–3,870 m), P. multiflora
(2,700–4,000 m), and P. elliptica (2,500–3,700 m).
Ecuadorian Pterichis are distributed along the Andes in three various ecoregions
(Olson et al., 2001;Fig. 2) and often two or more species co-occur in the area of 25 × 25 km
(Fig. 3). The highest number of genus representatives (13) was found growing in the
Eastern Cordillera real montane forests ecoregion (NT0121). Northern Andean páramo
(NT1006) is a home to ten Pterichis species and seven taxa occur in Northwestern Andean
montane forests (NT0145).
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 3/45
Taxonomic treatment
Pterichis Lindl., Gen. Sp. Orchid. Pl.: 444. 1840; Generitype: Pterichis galeata Lindl.
Terrestrial, rarely epiphytic, cespitose plants. Roots tuberous, fleshy, clustered, usually
1.5–5.0 mm in diameter. Leaves basal, few, rosulate, erect, shortly petiolate, blade usually
narrow, lanceolate. Stem simple, loosely sheathed. Inflorescence loosely to subdensely
few-flowered, lateral. Flowers non-resupinate, medium-sized, with the lateral sepals and lip
uppermost, usually reddish, maroon, brownish, yellowish, or polychromatic, sometimes
with dark freckles or lines. Ovary pubescent-glandular. Sepals subequal, free. Petals
narrower than the dorsal sepal, free or adnate to the dorsal sepal. Lip sessile, very broad,
concave in the center, variously ornamented with papillae or knob-like appendages along
margins, more or less 3-lobed; lateral lobes broad, surrounding the gynostemium;
middle lobe relatively narrow, fleshy, recurved. Gynostemium short, erect, massive.
Column part rudimentary, much shorter than the anther, column foot missing. Anther
erect, ovate or oblong-cordate, blunt to acute, motile. Pollinia 4, oblong-ovoid, relatively
compact, soft and powdery in P. triloba. Caudiculae formed from the apical and elongate
parts of pollinia. Staminodes thin and membranous or thick, fleshy, wing-like to lobed,
joined with the filament and the style below the stigma base, or entirely free, enveloping the
anther. Stigma ventral, confluent, flat to slightly convex, or horizontal, deeply concave,
pocket-like in P. triloba. Rostellum erect, narrow, filiform, delicate, soft, or canaliculate
with relatively broad base, truncate at the apex after the removal of the pollinium.
Viscidium single, detachable, cellular, thick, small, rounded or hood-like, asymmetric.
Hamulus usually present, finger-like, short or prominent, very obscure. Figs. 4 and 5.
The genus embraces about 45 species distributed from Costa Rica and Jamaica to
Bolivia. The majority of them are known from the Andes.
Figure 1 Altitudinal ranges of Ecuadorian Pterichis.Chart prepared in Microsoft Excel.
Full-size
DOI: 10.7717/peerj.10807/fig-1
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Figure 2 Terrestrial ecoregions of Ecuador with location of Pterichis populations. Ecoregion codes:
NT0107—Caqueta moist forests, NT0109—Cauca Valley montane forests, NT0115—Chocó-Darién
moist forests, NT0118—Cordillera Oriental montane forests, NT0121—Eastern Cordillera real montane
forests, NT0128—Iquitos várzea, NT0136—Magdalena Valley montane forests, NT0142—Napo moist
forests, NT0145—Northwestern Andean montane forests, NT0153—Peruvian Yungas, NT0156—Purus
várzea, NT0163—Solimões-Japurá moist forests, NT0166—Southwest Amazon moist forests, NT0174—
Ucayali moist forests, NT0178—Western Ecuador moist forests, NT0207—Cauca Valley dry forests,
NT0214—Ecuadorian dry forests, NT0221—Magdalena Valley dry forests, NT0223—Marañón dry
forests, NT0225—Patía Valley dry forests, NT0232—Tumbes-Piura dry forests, NT0905—Guayaquil
flooded grasslands, NT1004—Cordillera Central páramo, NT1006—Northern Andean páramo, NT1315
—Sechura desert, NT1405—South American Pacific mangroves. Map prepared in ArcGIS 9.3 (Esri,
Redlands, CA, USA). Full-size
DOI: 10.7717/peerj.10807/fig-2
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Key to the sections
1. Petals free from the dorsal sepal ...................................................................... sect. Pterichis
1Petals adnate to the dorsal sepal ........................................................................ sect. Acraea
Pterichis section Pterichis
Petals free from the dorsal sepal.
Figure 3 Richness of Pterichis species in Ecuador. Map prepared in ArcGIS 9.3 (Esri, Redlands, CA,
USA) using outcomes of species richness analysis conducted in DIVA-GIS.
Full-size
DOI: 10.7717/peerj.10807/fig-3
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Key to section section Pterichis
1. Petals almost linear, about 10 times longer than wide, sepals basally ovate, with margins
rolled up forming acuminate-caudate projection ............................................ P. aragogiana
1Petals ovate, elliptic to oblong-lanceolate, up to 5 times longer than wide, sepals apices
obtuse to acuminate, but not as described above .................................................................... 2
2. Floral bracts glabrous ................................................................................................................. 3
2Floral bracts glandular or ciliate ............................................................................................. 4
3. Petals glabrous, lip middle lobe constituting ca. 1/4 of the lip length ............. P. diuris
3Petals ciliate, lip middle lobe almost as long as the lip lamina ............. P. ansaloniana
4. Lip basal part without auricles or with inconspicuous auricles ....................................... 5
4Lip basal part with prominent auricles reaching almost the lip base ............ P. galeata
5. Lip basal part reniform .............................................................................................................. 6
5Lip basal part transversely elliptic ......................................................................... ; P. meirax
6. Petals subequal in length to dorsal sepal, lip wider than long .................. P. acuminata
6Petals longer than dorsal sepal, lip equally long and wide .......................... P.madsenii
Figure 4 Ecuadorian Pterichis sect. Pterichis (A–C) and sect. Acraea (D and E). (A–C) P. acuminata.
(D) P. multiflora. (E) P. triloba. Photos: M. Kolanowska. Full-size
DOI: 10.7717/peerj.10807/fig-4
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Figure 5 Habit of Ecuadorian Pterichis sect. Pterichis (A–G) and sect. Acraea (H–P). (A) Pterichis
acuminata—Dodson & Chase 17200 (RPSC). (B) Pterichis aragogiana—Holm-Nielsen et al. 29519 (AAU).
(C) Pterichis diuris—Funck & Schlim 1218 (W). (D) Pterichis galeata—Ulloa et al. 92355 (AAU).
(E) Pterichis meirax—Jameson s.n. (W). (F) Pterichis ansaloniana—Ulloa et al. 1361 (HA). (G) Pterichis
madsenii—Madsen & Ellemann 75293 (AAU). (H) Pterichis elliptica—528. (I) Pterichis habenarioides—
Huttel 528 (QCNE). (J) Pterichis multiflora—Løjtnant & Molau 13910 (AAU). (K) Pterichis parvifolia—
Álvarez et al. 2613 (QCNE). (L) Pterichis pauciflora—Minga & Verdugo 2530 (HA). (M) Pterichis sele-
niglossa—Dodson et al. 16396 (QCNE). (N) Pterichis triloba—Dodson et al. 10837 (Q). (O) Pterichis
tunguraguona—Holm-Nielsen et al. 5108 (AAU). (Q) Pterichis hirtziana—van der Werff & Palacios 8956
(QCNE). (P) Pterichis dodsoniana—Holm-Nielsen et al. 4767 (AAU).
Full-size
DOI: 10.7717/peerj.10807/fig-5
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Pterichis acuminata Schltr., Repert. Spec. Nov. Regni Veg. Beih. 7: 56. 1920. TYPE
(Garay, 1978: 182): Colombia. Antioquia. Alt. 3,200 m. M. Madero 27 (B†; Lectotype:
AMES-00103701!—drawing).
Plants 23–62 cm tall. Leaf, when present, petiolate; petiole 3.0–10.0 cm long; blade
5.5–10.0 × 2.5–4.0 cm, elliptic, acute. Scape glandular-ciliate in the upper part, glabrous at
base, enclosed by 4–5 tubular sheaths. Inflorescence 4.0–22.0 cm long, 5–20-flowered,
rachis densely ciliate. Flowers with pale green and faintly purplish tinted sepals which are
yellowish tinted with brown inside, dirty pale brownish purple petals, and reddish-purple
lip with darker brownish purple veins and white hairs. Floral bracts 8.0–12.3 mm long,
lanceolate to ovate-lanceolate, acute, externally glandular-ciliate. Pedicellate ovary
8.5–14.0 mm long, densely glandular. Dorsal sepal 7.9–9.8 × 2.5–4.2 mm, oblong-ovate to
ovate-ligulate, subacute, 3-veined, externally densely ciliate. Petals 7.5–10.0 × 1.8–4.0 mm,
free from dorsal sepal, obliquely oblong-lanceolate to lanceolate-sagittate, subobtuse,
3-veined, glabrous. Lateral sepals 8.8–10.0 × 2.2–3.6 mm, oblong-ovate to elliptic-
lanceolate, subacute to subobtuse, 3-veined, externally densely ciliate. Lip 5.6–7.4 ×
7.5–8.5 mm, obscurely 3-lobed, basal part reniform above truncate base; middle lobe
narrowly triangular to ovate, subobtuse, somewhat fleshy, recurved; disc papillate,
with irregular, knob-like projections along margins. Gynostemium 2.7–4.0 mm long.
Figs. 5 and 6.
Habitat and ecology: Terrestrial plants growing on roadsides, in cloud forest, humid
montane forest and paramo at the altitudes of 2,300–3,870 m. Flowering occurs in January,
April, June, and July.
Notes: This species is considered by some authors (Schweinfurth, 1941,1958) as conspecific
with Pterichis galeata, which is easily distinguished from P. acuminata by long-clawed
petals being longer than dorsal sepal (by 10–20%) and auriculate lip basal part.
In somewhat similar P. meirax the petals are oblong-lanceolate, with some hairs on the
margin and the lip basal part is transversely elliptic.
Representative specimens: Ecuador. Prov. Azuay. Parque Nacional Cajas. Km 33
Cuenca-Molleturo. Sendero alrededor de Laguna Cucheros. Alt. 3,820–3,870 m.
17 January 2003. C. Ulloa et al. 1260 (HA!). Prov. Morona-Santiago. Cantón Gualaquiza.
Area de Bosque Vegetación Protectora Tambillo. Alt. 2,900 m. 26 June 2001. L. Suin &
J. Guartán 1014 (HA!, QCNE!). Prov. Morona-Santiago. Between San José de Raranga and
San Miguel de Cuyes. Alt. ca 2900 m. 9 July 2017. M. Kolanowska et al. E17/73 (photo!).
Prov. Napo. Road Quito to Baeza, above Papallacta. On lava flow at Lago Papallacta.
Alt. 3,100 m. 22 Jun 1987. C.H. Dodson & M. Chase 17200 (RPSC!); Summit and
uppermost N slopes of Cerro Sumaco. Virgin moist páramo. Alt. 3,700–3,840 m. 26 April
1979. B. Løjtnant & U. Molau 12786 (AAU!); km 30 on road from El Carmelo towards La
Bonita, SE of Santa Bárbara. Alt. ca. 2,600 m. 13 April 1979. B. Løjtnant et al. 12402
(AAU!); E-facing ridge on the N side of Cerro Sumaco. Virgin moist páramo with scattered
shrubs, on steep slope. Alt. 3,400–3,600 m. 25 April 1979. B. Løjtnant & U. Molau
12744 (AAU!). Prov. Sucumbíos. El Mirador, Playon de San Francisco-Julio Andrade,
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km 12. Alt. 3,200–3,400 m. 11 Jul 1991. C.H. Dodson, N. Williams & M. Whitten 18781
(RPSC!, UGDA-DLSz!—drawing).
Pterichis cf. acuminata
In the following collections the general plant and flower appearance correspond to
P. acuminata, but some details of the examined specimens distinguish them from the
typical form of this species:
Figure 6 Pterichis acuminata Schltr. (A) Floral bract. (B) Dorsal sepal. (C) Lateral sepal. (D) Petal.
(E) Lip. Scale bars = 1 mm. Drawn by S. Nowak from Suin & Guartán 1014 (QCNE).
Full-size
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–Prov. Imbabura. Cotacachi Cantón. Parroquia Plaza Gutierrez. Table Chupa, arriba de
Apuela. Bosque húmedo montano. Alt. 2,300–3,000 m. 12 May 1992. G. Tipaz et al. 1000
(QCNE!)—cf. P. acuminata but with prominent, truncate lip basal part
–Prov. Morona-Santiago. Road Gualaceo-El Limon. From the pass towards El Limon.
Paramo or scrub just below paramo. Alt. 3,200–3,400 m. 8 February 1989. H. van der
Werff & W. Palacios 10516 (QCNE!)—cf. P. acuminata but with prominent, truncate lip
basal part
–Prov. Carchi [Sucumbíos]. Road Tulcán-El Carmelo, at km 10. Roadbank in patch of
montane shrub in bunchgrass paramo. Alt. 3,350 m. 27 May 1980. H. Balslev &
F. Quintana 23834 (AAU!)—cf. P. acuminata but petal elliptic, 1-veined
–Prov. Sucumbíos. Alt. ca. 3,300 m. 6 July 2017. M. Kolanowska E17/38 (photo!)—cf.
P. acuminata but with petals microscopically, sparsely ciliate along margin
–Prov. Chimborazo. Next to the road Toncal de la Sierra. 8 July 2017. Alt. 3,200 m.
M. Kolanowska et al. E17/51(photo!)—cf. P. acuminata but with prominent, truncate lip
basal part and petals microscopically, sparsely ciliate along margin.
Pterichis aragogiana Szlach. & Kolan., Ann. Bot. Fenn. 56(1–3): 176–178. 2019.
TYPE: Ecuador. Prov. Azuay. Páramo de Matanga, km 25 on road Sigsig–Gualaquiza
(old muletrack), W of the pass. Alt. 3,150 m. 14 December 1980. L.B. Holm-Nielsen et al.
29519 (holotype AAU!).
Plants 40–55 cm tall, leafless during flowering. Scape glandular in upper third,
enveloped in 7 tubular, acute sheaths of which upper ones are glandular-ciliate.
Inflorescence ca. 8.5 cm long, densely ciliate. Flowers externally olive-green, petals and
internal part of sepals orange-red, lip orange red flushed with violet in basal part of lip
middle lobe, sepals externally densely glandular-ciliate. Floral bracts 14.0–20.0 mm long,
ovate, cucullate, externally densely ciliate. Pedicellate ovary ca. 13.0 mm long, densely
glandular-ciliate. Dorsal sepal 11.0–14.5 × 2.8–4.5 mm, ovate in lower half, apical part
narrow, rolled up along margins hence appearing acuminate-caudate, subobtuse, 3-veined.
Petals 11.0–14.5 × 1.0–1.4 mm, free from dorsal sepal, linear, subobtuse, 3-veined,
glabrous. Lateral sepals 9.5–15 × 3–3.5 mm, obliquely ovate in lower half, rolled up
along margins above giving an acuminate-caudate appearance, subobtuse, 3-veined.
Lip 9.0–12.0 × 10.0–11.0 mm, hastate, basal part reniform above truncate base; middle lobe
ca half of the lip length, ovate, acuminate; disc glandular, primarily 9-veined, lateral veins
branching, ornamented with knob-like projections along margins. Gynostemium
3.0–5.8 mm long. Figs. 5 and 7.
Habitat and ecology: Terrestrial plants growing in scrub páramo with Blechnum L.
(Blechnaceae), Befaria Mutis (Ericaceae), Osteomeles Lindl. (Rosaceae), and Cortaderia
Stapf. (Poaceae) at the altitude of 3,150 m. Flowering in October and December.
Notes: Pterichis aragogiana is easily distinguished from all other genus representatives by
the caudate sepals. Their lower part is more or less ovate and the upper one is rolled up
along margins forming a caudate, acuminate projection. Unlike in other species of the
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nominal section of Pterichis, the lip middle lobe of P. aragogiana constitutes ca. half of the
total lip length.
Representative specimens: Ecuador. Prov. Azuay. Páramo de Matanga, km 25 on road
Sigsig–Gualaquiza (old muletrack), W of the pass. Alt. 3,150 m. 14 December 1980.
Figure 7 Pterichis aragogiana Szlach. & Kolan. (A) Dorsal sepal. (B) Lateral sepal. (C) Petal. (D) Lip.
Scale bars = 1 mm. Drawn by N. Olędrzy
nska from Holm-Nielsen & al. 29519 (AAU).
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L.B. Holm-Nielsen et al. 29519 (AAU!). Prov. Morona-Santiago. Road from Sigsig to
El Pangui. Páramo Matanga. 311′49″S, 7846′08″W, 4 October 2003. M. Blanco et al.
2515 (FLAS!). Prov. Zamora Chinchipe. Cerro Plateado. Á.J. Pérez 1355 (QCA!).
Pterichis diuris Rchb. f., Bonplandia (Hannover) 2: 10. 1854. TYPE: (Kolanowska &
Olędrzy
nska, 2015: 44): Venezuela. Mérida. N. Funck & L.J. Schlim 1218 (lectotype W-R
377!, isolectotype AMES-00083567—microscope slide; AMES-00103705!—drawing)
Plants 40 cm long. Leaf up to 15.0 cm long, petiolate, petiole up to 7.0 cm long; blade
5.8–7.6 × 2.3 cm, oblong-elliptic, obtuse. Scape with 4–5 tubular, glabrous sheaths.
Inflorescence 7.0–8.0 cm long, ciliate, subdensely several-flowered. Floral bracts 13.0 mm
long, ovate, obtuse, glabrous. Pedicellate ovary 14.0 mm long, pubescent. Flowers with
dirty green sepals, translucent, brownish-yellow petals and yellow, brown-veined lip.
Dorsal sepal 8.3 × 3.0 mm, ovate-lanceolate, obtuse, 3- or 5-veined, externally ciliate. Petals
10.2 × 2.0 mm, free from dorsal sepal, obliquely linear-lanceolate, obtuse, 3-veined,
glabrous. Lateral sepals 9.0 × 4.5 mm, obliquely ovate, subacute, 3- or 5-veined, externally
sparsely ciliate. Lip 6.1 ×10.0 mm, basal part reniform in outline with lateral auricles, base
truncate; middle lobe about 1/4 of the total lip length, narrowly triangular-lanceolate,
papillate; disc ornamented with large glands along the margin, primarily 5-veined, lateral
veins branching. Gynostemium 3.0 mm long. Figs. 5 and 8.
Habitat and ecology: Terrestrial plants growing in montane forest, in steep creek with large
ferns at the altitude of 3,100–3,150 m. Flowering occurs in April.
Notes: From all other Ecuadorian species of nominal section of Pterichis, P. diuris differs in
having glabrous floral bracts. In the presence of the auricles, the lip of this taxon resembles
somewhat P. galeata which differs from P. diuris by obliquely elliptic petals which are
not long-clawed.
Representative specimen: Ecuador. Prov. Napo. N side of Cerro Sumaco, loma NW of
campsite. Alt. 3,100–3,150 m. 28 April 1979. L.B. Holm-Nielsen et al. 17385 (AAU!).
Pterichis galeata Lindl., Gen. Sp. Orchid. Pl. 445. 1840. TYPE (Kolanowska & Szlachetko,
2017: 91): Peru. A. Mathews s.n. (lectotype K-L-000079985!).
Plant 25–30 cm tall. Leaf (when present) petiolate; petiole 2.0 cm long; blade 4.0 ×
1.0 cm, oblong-elliptic, obtuse. Scape glandular-pilose above, provided with 4–5, more or
less distant, tubular sheaths, the lower 2 glabrous, the upper ones pilose. Inflorescence
4.0–8.0 cm long, sublaxly 5–7-flowered, rachis densely ciliate. Flowers greenish yellow with
brown veins. Floral bracts ca. 11 mm, ovate to lanceolate, ciliate. Pedicellate ovary 13.0 mm
long, densely ciliate. Dorsal sepal 9.2 × 3.0 mm, ovate-lanceolate, obtuse, 3-veined,
externally ciliate. Lateral sepals 9.3 × 2.6 mm, obliquely lanceolate to lanceolate-ovate,
acuminate, 3-veined, externally ciliate. Petals 10.0 × 3.3 mm, free from dorsal sepal, obliquely
lanceolate-elliptic, unguiculate or narrowed below, obtuse, 3-veined, lateral veins branching,
glabrous. Lip 7.1 × 8.3 mm, obscurely 3-lobed, basal part broadly reniform- with pair
of auricles on edges; middle lobe triangular or triangular-lanceolate, densely hairy;
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 13/45
disc papillate with series of irregular glands along margin. Gynostemium 3.0 mm long.
Figs. 5 and 9.
Habitat and ecology: Terrestrial on roadside and in secondary vegetation dominated by
Freziera Willd. (Theaceae) at the altitude of 3,050–3,100 m. Flowering in Ecuador occurs
in August.
Notes: This species can be distinguished from other Ecuadorian representatives of Pterichis
sect. Pterichis by having long-unguiculate petals. In examined Ecuadorian specimens the
petal claw is shorter than in typical form of P. galeata but without additional material
we are not able to define them as a new species.
Representative specimen: Ecuador. Prov. Carchi. Julio Andrade-El Carmelo, turn off towards
El Ajún, km 0–3. Alt. 3,050–3,100 m. 10 August 1990. C. Ulloa et al. 92355 (AAU!).
Figure 8 Pterichis diuris Rchb. f. (A) Dorsal sepal. (B) Lateral sepal. (C) Petal. (D) Lip. Scale
bars = 1 mm. Drawn by S. Nowak from Holm-Nielsen & al. 17385 (AAU).
Full-size
DOI: 10.7717/peerj.10807/fig-8
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 14/45
Pterichis meirax Rchb. f. ex Szlach. & Kolan., Ann. Bot. Fenn. 51(5): 331. 2014. TYPE:
Ecuador. W. Jameson s.n. (holotype W 870!).
Plants 10–22 cm tall. Leaf petiolate; petiole 2.0–8.0 cm long; blade 3.5–13.0 ×
0.9–1.4 cm, lanceolate to oblong-lanceolate, acute. Scape ciliate above the basal third, with
1 glabrous and 4 ciliate sheaths. Inflorescence 3.5–7.0 cm long, laxly 5–9-flowered,
rachis densely ciliate. Flowers with dirty brownish-green sepals, dirty brownish purple
petals tipped with yellow, and reddish to reddish-brown lip tipped with yellow and with
white hairs. Floral bracts 6–17 mm long, externally ciliate. Pedicellate ovary 7–20 mm long,
densely ciliate. Dorsal sepal 6.0–8.3 × 2.0–3.3 mm, ovate-lanceolate, obtuse, 3-veined,
externally glandular to ciliate. Petals 7.0–8.0 × 2.3–3.0 mm, free from dorsal sepal,
Figure 9 Pterichis galeata Lindl. (A) Floral bract. (B) Dorsal sepal. (C) Lateral sepal. (D) Petal. (E) Lip.
Scale bars = 1 mm. Drawn by S. Nowak from Ulloa & al. 92355 (AAU).
Full-size
DOI: 10.7717/peerj.10807/fig-9
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 15/45
obliquely oblong-lanceolate to linear-lanceolate, subobtuse, glabrous, 3-nerved. Lateral
sepals 6–8.2 × 2.5–3.2 mm, obliquely ovate, concave, acute, 3-veined, externally glandular
to ciliate. Lip 5.5–6.3 × 8.0–9.0 mm, basal part transversely elliptic in outline above
truncate base; middle lobe ca 1/3 of the lip length, ovate, papillate; disc papillate with
numerous knob-like projections along margin in the upper half, 7- or 9-veins with lateral
veins branching. Gynostemium 3.0–3.6 mm long. Figs. 5 and 10.
Habitat and ecology: Terrestrial in elfin forest and steep slopes along road at the altitude of
3,200–3,500 m. Flowering occurs in April and May.
Notes: This species resembles Pterichis acuminata from which it differs by the petals shape
(oblong-lanceolate in P. meirax) and transversely elliptic lip basal part (reniform in
P. acuminata). P. meirax resembles P. habenarioides from which it differs mainly by its lip
form. The lip of P. habenarioides is triangular-elliptic in outline, with subtriangular apical
Figure 10 Pterichis meirax Rchb. f. ex Szlach. & Kolan. (A) Floral bract. (B) Dorsal sepal. (C) Lateral
sepal. (D) Petal. (E) Lip. Scale bars = 1 mm. Drawn by S. Nowak from Løjtnant & al. 13755 (AAU).
Full-size
DOI: 10.7717/peerj.10807/fig-10
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 16/45
part, pubescent margins and interior papillae. The papillae on P. meirax lip are distributed
also along the thickened apical margins below the ligulate apex.
Representative specimens: Ecuador. Prov. Carchi. Carmelo, road from Tulcán to Santa
Barbara. Alt. 3,200 m. May 1985. A. Hirtz 2617 (RPSC!, UGDA-DLSz!—drawing). Prov.
Cotopaxi. Latacunga-Quevedo road, above Pilaló, km 74 from Pujili. Alt. 3,500 m.
26 April 1979. B. Løjtnant et al. 13755 (AAU!). Prov. Tungurahua. W. Jameson s.n. (W!).
Pterichis ansaloniana Kolan., Szlach. & S. Nowak, sp. nov.TYPE: Ecuador. Prov. Azuay.
Parque Nacional Cajas. Km 35.7 Cuenca-Molleturo. Sendero Tres Cruces. Laguna
Negra-Laguna Larga. Páramo de pajonal. Alt. 4,100 m. 23 January 2003. C. Ulloa et al. 1361
(holotype HA 5333!—left hand plant; isotype HA 5333!).
Species similar to Pterichis diuris distinguished by more or less lanceolate, ciliate petals,
broadly truncate lip base, presence of prominent knob-like projections on the lip margins,
and larger lip middle lobe.
Plant 18–30 cm tall. Leaf unknown. Scape with 5–6 prominent sheaths. Inflorescence ca.
6 cm long, congested, few-flowered, rachis ciliate. Floral bracts 10 mm long, ovate,
rounded, glabrous. Pedicellate ovary 13.5 mm long, sparsely ciliate. Flowers with green
sepals and yellow lip with brown-purple lines. Dorsal sepal 6.0 × 2.4 mm, oblong ovate,
rounded at the apex, 3-veined, externally sparsely ciliate. Petals 8.9 × 1.8 mm, free from
dorsal sepal, obliquely lanceolate, widened in the basal third, 3-veined, sparsely ciliate.
Lateral sepals 6.5 × 2.7 mm, obliquely ovate, acuminate, obtuse, 3-veined, externally
sparsely ciliate. Lip 5.5 × 7 mm, reniform above truncate base; middle lobe ca half of the lip
length, elliptic, apiculate, papillate; disc papillate with series of knob-like projections along
the margin. Gynostemium 2.8 mm long. Figs. 5 and 11.
Etymology: Dedicated to Raffaella Ansaloni, the curator of Herbario Azuay.
Habitat and ecology: Terrestrial in paramo at the altitude of 4,100 m. Flowering in January.
Notes: This species resembles Pterichis diuris from which it can be distinguished by more
or less lanceolate, ciliate petals (vs. glabrous, obliquely elliptic), broadly truncate lip base,
presence of prominent knob-like projections on the lip margins (vs. with large glands
along the margin), and larger lip middle lobe (almost as long as the lip lamina vs.
constituting ca. 1/6 of the lip length). The similar, long lip middle lobe is observed in
P. aragogiana, but it this species sepals are caudate and petals are linear and glabrous.
Type collection deposited in HA consists of two plant, and the holotype is plant on the
left hand with mature inflorescence.
Representative specimen: Ecuador. Prov. Azuay. Parque Nacional Cajas. km 35.7 Cuenca-
Molleturo. Sendero Tres Cruces. Laguna Negra-Laguna Larga. Páramo de pajonal. Alt.
4,100 m. 23 January 2003. C. Ulloa et al. 1361 (HA!).
Pterichis madsenii Kolan., Szlach. & S. Nowak, sp. nov.TYPE: Ecuador. Prov. Loja. Parque
Nacional Podocarpus. Above Nudo de Cajanuma, trail to Mirador. Scrub and ridge-top
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 17/45
vegetation above tree limit. Alt. 3,000–3,150 m. 6 September 1988. J.E. Madsen &
L. Ellemann 75293 (holotype AAU!).
Species similar to Pterichis acuminata but with sparsely ciliate sepals, prominent lip
middle lobe and lip being as long as wide.
Plants ca. 70 cm tall. Leaf petiolate; petiole 12.0 cm long; blade 13.0 × 3.0 cm, narrowly
elliptic, acute. Scape glandular-ciliate above basal part, enclosed by 8 tubular sheaths.
Inflorescence 9.0 cm long, ca. 10-flowered, rachis densely ciliate. Flower color not
recorded. Floral bracts ca. 11.0 mm long, ovate, acute, externally glandular-ciliate.
Pedicellate ovary 14.0 mm long, pubescent. Dorsal sepal 8.0 × 3.0 mm, oblong-ovate,
obtuse, 3-veined, externally sparsely ciliate. Petals 9.0 × 3.8 mm, free from dorsal sepal,
obliquely elliptic, sub-bilobulate, rounded, 3-veined, glabrous. Lateral sepals 9.2 × 3 mm,
oblong-ovate, subobtuse, 3-veined, externally sparsely ciliate. Lip 7.0 × 7.1 mm, basal part
reniform, subauriculate; middle lobe ca half of the lip length, triangular, obtuse, fleshy;
Figure 11 Pterichis ansaloniana Kolan., Szlach. & S. Nowak, sp. nov.(A) Floral bract. (B) Dorsal sepal.
(C) Lateral sepal. (D) Petal. (E) Lip. Scale bars = 1 mm. Drawn by S. Nowak from Ulloa & al. 1361
(HA). Full-size
DOI: 10.7717/peerj.10807/fig-11
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 18/45
disc papillate, with irregular, knob-like projections along margins, 7-veined, veins apically
branching. Gynostemium 3.0 mm long. Figs. 5 and 12.
Etymology: Dedicated to Jens E. Madsen, senior collector of the type specimen.
Habitat and ecology: Terrestrial in scrub and ridge-top vegetation above tree limit at the
altitudes of 3,000–3,150 m. Flowering in September.
Notes: Species similar to Pterichis acuminata but with sparsely ciliate sepals (vs. sepals
densely ciliate), prominent lip middle lobe constituting half of the total lip length (vs. up to
1/3 of the total lip length) and lip being as long as wide (vs. wider than long).
Figure 12 Pterichis madsenii Kolan., Szlach. & S. Nowak, sp. nov.(A) Floral bract. (B) Dorsal sepal.
(C) Lateral sepal. (D) Petal. (E) Lip. Scale bars = 1 mm. Drawn by S. Nowak from Madsen & Ellemann
75293 (AAU). Full-size
DOI: 10.7717/peerj.10807/fig-12
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 19/45
Representative specimen: Ecuador. Prov. Loja. Parque Nacional Podocarpus. Above Nudo
de Cajanuma, trail to Mirador. Alt. 3,000–3,150 m. 6 September 1988. J.E. Madsen &
L. Ellemann 75293 (AAU!).
Pterichis section Acraea (Lindl.) Kolan. & Szlach.
Ann. Missouri Bot. Gard. 102(1): 97. 2017. ≡Acraea Lindl., Pl. Hartw.: 155. 1845; Type:
Acraea parvifolia Lindl., Pl. Hartw.: 155. 1845.
Petals adnate to the dorsal sepal.
Key to section Section Acraea
1. Petals glabrous ...............................................................................................................................2
1Petals ciliate ................................................................................................................................. 6
2. Truncate basal part of the lip constituting less than 1/3 of the lip lamina ................... 3
2Truncate basal part of the lip constituting about 1/2 of the lip lamina ........................ 5
3. Petals narrowly ovate above narrow claw ................................................ P. tunguraguona
3Petals linear-oblong to almost filiform ................................................................................. 4
4. Floral bracts densely glandular-pubescent ......................... P. multiflora var. multiflora
4.Floral bracts glabrous ........................................................ P. multiflora var. ecuadorensis
5. Petals 1-veined ..................................................................................................... P. seleniglossa
5Petals 3-veined ............................................................................................................. P. triloba
6. Lip basal part transversely elliptic, widest near the middle .............................................. 7
6Lip basal part more or less reniform or lunate, widest below the middle .................... 8
7. Lip middle lobe broadly ovate-ligulate, constitutes less than 1/3 of the total lip length
.............................................................................................................. P. elliptica var. elliptica
7.Lip middle lobe oblong-elliptic, constitutes more than 1/3 of the total lip length ...............
......................................................................................................................... P. elliptica var. ecuadorensis
8. Lip lamina lunate .................................................................................................... P. hirtziana
8Lip lamina reniform .................................................................................................................. 9
9. Lip triangular in outline with middle lobe not well-separated .................. P. pauciflora
9Lip with distinctly separated middle lobe .......................................................................... 10
10. Lip lacking knob-like projections along the margin ....................................................... 11
10Lip with knob-like projections along the margin ........................................ P. parvifolia
11. Petals 1.2 mm wide, linear, lip middle lobe constituting over 1/3 of the total lip length
...................................................................................................................................... P. dodsoniana
11Petals 1.5–2.5 mm wide, obliquely lanceolate, lip middle lobe constituting not more
than 1/4 of total lip length ............................................................................... P. habenarioides
Pterichis elliptica Kolan. & Szlach., Wulfenia 22: 222. 2015. TYPE: Ecuador. Zamora-
Chinchipe, Road from Loja to Zamora, km 14. Alt. 2,800 m. 18 November 1961.
C.H. Dodson & L.B. Thien 1326 (holotype RPSC 0000936-H!; isotype: RPSC 0000936-I!).
Plants (12)27–40 cm tall. Leaf petiolate; petiole 5.0–9.0 cm long, blade up to 6.0 ×
2.0 cm, elliptic, acute. Scape sparsely glandular at the base, densely near the apex, with
4–5 sheaths. Inflorescence 3.0-8.0 cm long, 3–9-flowered, rachis densely ciliate. Flowers
greenish yellow or brownish yellow with maroon stripes. Floral bracts 7.2–11.8 mm long,
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 20/45
elliptic-lanceolate to elliptic, obtuse to acuminate, glabrous or almost so. Pedicellate ovary
11.0–18.0 mm long, densely glandular-ciliate. Dorsal sepal 5.5–8.3 × 1.6–2.5 mm, ovate to
ovate-lanceolate, obtuse, 3-veined, externally almost glabrous, glandular-ciliate near the base.
Petals 5.2–10.0 × 1.5–2.2 mm, agglutinate to dorsal sepal, obliquely oblong-lanceolate,
obtuse, 3-veined, sparsely ciliate. Lateral sepals 6–8×3–3.8 mm, obliquely ovate, obtuse,
apically canaliculated, 3- or 4-veined, externally ciliate. Lip 5.3–7.0 × 7.5–8.8 mm, basal part
transversely elliptic; middle lobe ca 1/3 of the lip length, broadly ovate-ligulate, papillate; disc
papillate, with numerous knob-like projections along the margins, 9- or 11-veined.
Gynostemium 2.3–3.2 mm long. Figs. 5 and 13.
Figure 13 Pterichis elliptica Kolan. & Szlach. (A) Floral bract. (B) Dorsal sepal. (C) Lateral sepal.
(D) Petal. (E) Lip. Scale bars = 1 mm. Drawn by S. Nowak from Huttel 528 (QCNE).
Full-size
DOI: 10.7717/peerj.10807/fig-13
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 21/45
Habitat and ecology: Terrestrial plant growing in trail embankment, montane forest and
wet páramo vegetation with numerous shrubs or large patches of Neurolepis Meins.
(Poaceae) or with Blechnum and thick layers of mosses at the altitude of 2,500–3,700 m.
Flowering occurs in January, April, September, October, November and December
Notes: Pterichis elliptica resembles P. parvifolia from which it differs by the elliptic basal lip
part (vs. reniform), rounded lip base (vs. truncate), and lip middle lobe constituting ca. 1/3
of the lip length (vs. middle lobe shorter than 1/2 of the total lip length).
Representative specimens: Ecuador. Prov. Azuay. Road Sigsig-Gualaquiza, km 25.6. At the
pass on military post road. Wet páramo vegetation with large patches of Neurolepis.
Km 3.3. from pass to military post. Alt. 3,200–3,330 m. 11 January 2000. P.M. Jørgensen
et al. 1810 (QCNE!). Prov. Loja. Along path (camino de herradura) at pass at km 13.5
Loja-Zamora. Alt. 2,800 m. 21 Sep 1980. C.H. Dodson, C. Luer, J. Luer, P. Morgan,
H. Morgan, A. Perry & J. Kuhn 10519 (RPSC!); Parque Nacional Podocarpus. Cerro
Toledo. Montane forest and paramo. Alt. 2,500–3,400 m. 30 October 1989. J.E. Madsen
86289 (AAU!, LOJA!, QCA!). Prov. Morona-Santiago. Sigsig-Gualaquiza, km 26,
turnoff towards military antenas. Páramo with many shrubs. Alt. 3,290–3,570 m.
2 December 1990. P.M. Jørgensen et al. 92777 (AAU!, QCA!). Prov. Napo. N side of Cerro
Sumaco. 100 m NW of campsite. Alt. 3,700 m. 24 April 1979. L.B. Holm-Nielsen et al.
17128 (AAU!). Prov. Pichincha. Papallacta. Alt. 3,600 m. 27 December 1984. Ch. Huttel
528 (QCNE!). Prov. Zamora-Chinchipe. Road from Loja to Zamora, km 14. Alt. 2,800 m.
18 November 1961. C.H. Dodson & L.B. Thien 1326 (RPSC 0000936-H!, RPSC
0000936-I!).
The below listed collection could be classified in P. elliptica but it is characterized by a
very long lip middle lobe, not observed in other populations. Moreover, it was found
growing epiphytically on Polylepis Ruiz & Pav. (Rosaceae) in humid montane forest while
Pterichis almost always are terrestrial plants. In our opinion these features are sufficient to
consider that as separated variety.
Pterichis elliptica var. ecuadorensis Kolan., Szlach. & S. Nowak, var. nov.TYPE: Ecuador.
Prov. Napo. Cantón Quijos. Parroquia Papallacta, sector Río Papallacta. Alt. 3,400 m.
2 November 1997. Flowers green. S. Teran & P. Quela 26 (holotype: QAP 33285!). Fig. 14.
Distinguished from typical form of Pterichis elliptica by the epiphytic habit, by the long,
oblong-elliptic lip middle lobe which constitutes more than 1/3 of the total lip length.
Etymology: In reference to the country of origin of the type collection.
Pterichis habenarioides (F. Lehm. & Kraenzl.) Schltr., Repert. Spec. Nov. Regni Veg. Beih.
7: 214. 1920. ≡Goodyera habenarioides F. Lehm. & Kraenzl., Bot. Jahrb. Syst. 26: 499. 1899.
TYPE (Garay, 1978: 183): Colombia. Cauca. Paramo Guanacas. F. Lehmann 6419
(lectotype K-000881685!).
Pterichis costaricensis Ames & C. Schweinf., Sched. Orch. 10: 10. 1930. TYPE
(Garay, 1978: 183): Costa Rica. H. E. Stork 2344 (lectotype: AMES-00103704!,
isolectotypes: AMES-00083566!—microscope slide, US; UGDA-DLSz!—drawing).
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 22/45
Plants 34 cm tall. Leaf usually petiolate; petiole up to 12.0 cm long; blade 7.0–12.0 ×
1.0–2.0 cm, narrowly lanceolate, acuminate. Scape pubescent, remotely 4–5-sheathed,
sheaths glabrous. Inflorescence 7.0–10.0 cm long, sublaxly 10–12-flowered, rachis
densely ciliate. Flowers yellow to green, sepals brown-spotted, petals greenish tinted, lip
varying from deep green to yellow with dark purple or brownish markings. Floral bracts
8.3–9.0 mm long, ovate-lanceolate to ovate, obtuse, glabrous or only basally ciliate.
Pedicellate ovary 9.0–11.5 mm long, puberulent. Dorsal sepal up 6.7–9.0 × 2.5–3.0 mm,
ovate, obtuse, 3-veined, glabrous or externally sparsely ciliate. Petals 6.9–8.3 × 1.5–2.5 mm,
agglutinate to dorsal sepal, obliquely lanceolate, falcate, with ligulate terminal third, 2- or
3-veined, sparsely ciliate along margins. Lateral sepals 5.7–7.8 × 3–3.8 mm, obliquely
Figure 14 Pterichis elliptica var. ecuadorensis Kolan., Szlach. & S. Nowak, var. nov.(A) Floral bract.
(B) Dorsal sepal. (C) Lateral sepal. (D) Petal. (E) Lip. Scale bars = 1 mm. Drawn by S. Nowak from Teran
& Quela 26 (QAP). Full-size
DOI: 10.7717/peerj.10807/fig-14
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 23/45
broadly ovate, acuminate with involute margins, glabrous or externally sparsely ciliate.
Lip 5.3–7.0 × 8.0 mm, 3-lobed, basal part reniform with small auricles; middle lobe ca 1/3 f
the lip length, triangular-ovate, apiculate, pubescent; disc papillate, with numerus
glands along the margin, 5-veined with branching veins. Gynostemium 2.3–2.8 mm long.
Figs. 5 and 15.
Habitat and ecology: Terrestrial plant found in xerophytic scrub, intermingled with
meadows and dryer grassland at the altitude of 3,250 m and in paramo dominated by
Calamagrostis Adans. (Poaceae) and Espeletia pycnophylla Cuatrec. (Asteraceae) at
3,250–3,800 m. Flowering occurs in May and July.
Notes: Pterichis habenarioides resembles P. parvifolia from which it differs by the lip
margin being ornamented with numerus glands along the margin (vs. lip with knob-like
Figure 15 Pterichis habenarioides (F. Lehm. & Kraenzl.) Schltr. (A) Floral bract. (B) Dorsal sepal.
(C) Lateral sepal. (D) Petal. (E) Lip. Scale bars = 1 mm. Drawn by S. Nowak from Narváez 586
(QCNE). Full-size
DOI: 10.7717/peerj.10807/fig-15
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 24/45
projections in P. parvifolia). This species is also similar to Colombian endemic
P. colombiana from which it differs by the glabrous sheaths on the scape, as well as by the
petals form (falcate in the middle part) and the dorsal sepal size (much larger than the
lateral sepals; Morales, 1986). Pterichis habenarioides Schtr. differs from P. triangularilabia
Kolan. & Szlach. by transversely triangular-elliptic lip with triangular to ligulate apex.
Representative specimens: Ecuador. Prov. Azuay. Paramo de Tinajillas between Cuenca
and Oña. 27 July 1982. S.E. Clemants et al. 2200 (NY!, QCA!); Km 67 S of Cuenca on
Panamerican Highway. Alt. 3,250 m. 4 May 1973. L.B. Holm-Nielsen et al. 4985 (AAU!).
Prov. Carchi. San Pedro de Huaca. Estación Biológica Guandera. Parroquia Marsical Sucre.
Alt. 3,500 m. 12 July 2000. E. Narváez 586 (QCNE!). Prov. Imbabura. Timber line
vegetation on Hacienda Yura Cruz. 10 km N of Ibarra. Alt. 3,700–3,800 m. 25 May 1973.
L.B. Holm-Nielsen et al. 6492 (AAU!).
Pterichis multiflora (Lindl.) Schltr., Bot. Jahrb. Syst. 45: 389. 1911. Acraea multiflora
Lindl., Orchid. Linden.: 26. 1846. TYPE (Garay, 1978: 185): Venezuela. Mérida. J. Linden
673 (lectotype K-L, K—Garay’s illustration!, isolectotypes: G-00169178!, P?).
Plants 13–55 cm tall. Leaf petiolate; petiole 4.0–12.0 cm long; blade 7.0–14.0 cm long, up
to 3.5 cm wide, elliptic to oblong-lanceolate, subobtuse. Scape glandular-ciliate above
basal sheath or from the base, enclosed with 3–7 sheaths, the uppermost ciliate.
Inflorescence 3.0–18.0 cm long, subdensely 5–20-flowered, rachis densely ciliate. Flowers
with dirty yellowish-green or dirty brown-green sepals with darker greenish-brown veins,
petals brown or reddish brown in the basal third, pale yellowish-brown in the center
and yellowish at the apex, lip brownish yellow with dark brown veins. Floral bracts
7.1–11.0 mm long, ovate-lanceolate, acuminate, densely glandular-pubescent. Pedicellate
ovary 11.0–18.0 mm long, pubescent. Dorsal sepal 7.1–9.0 × 2.3–3.4 mm, ovate-lanceolate,
obtuse to acuminate, 3- or 5-veined, externally ciliate. Petals 7.2–10.1 × 1.2–2.3 mm,
agglutinate to dorsal sepal, linear-oblong to almost filiform, 2- or 3-veined, glabrous.
Lateral sepals 5.2–11.0 × 2.7–5.0 mm, oblique, ovate-lanceolate to ovate, obtuse to
acuminate, 3-veined, externally ciliate. Lip 4.2–8×7–10 mm, obscurely 3-lobed, basal part
reniform above truncate base; middle lobe ca ½ of the lip length, rather large in proportion,
ovate-triangular, recurved, densely papillose-hirsute; disc papillate with numerous
glands along margin. Gynostemium 2.1–3.5 mm long.
Habitat and ecology: Rupicolous, in grass páramo and virgin elfin forest as well as on slopes
along road in cloud forest at the altitudes of 2,700–3,850 m. Flowering occurs in January,
March, May, June, July, and August.
Notes: This species is similar to Pterichis triloba which, however, has oblong-elliptic to
oblong-lanceolate petals widened near the middle. The 2–3-veined petals makes
P. multiflora somewhat similar to P. triangularilabia and P. habenarioides but in
P. multiflora petals are 5–6 times longer than wide (vs. 3–4 times longer than wide).
The illustration of type specimen deposited in K was made based on plant deposited in the
same herbarium, however, we did not find this specimen among herbarium sheets.
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 25/45
Collection Linden 673 deposited in P herbarium (P00363510) represents not Pterichis
but Ponthieva.
Representative specimens: Ecuador. Prov. Azuay. Páramo de Tinajillas, km 23-28 from
Cumbe on road to Loja. Alt. 3,200–3,300 m. 16 June 1979. B. Løjtnant et al. 14951 (AAU!).
Prov. Cotopaxi. Latacunga-Quevedo road, 3–5 km above Pilaló. Alt. 2,700–2,800 m.
28 May 1979. B. Løjtnant & U. Molau 13910 (AAU!); Volcán Cotopaxi. Along the road to
and at Limpio Punga. Alt. 3,500–3,850 m. 23 March 1984. S. Laegaard 51851 (AAU!).
Prov. Morona-Santiago. Along road from Cuenca to Limon. Alt. 3,000 m. Jan 1989.
C. Luer, J. Luer, A. Andreetta, P. Jesup & S. Ortega sub A. Hirtz 4171 (RPSC!, UGDA-DLSz!
—drawing). Prov. Pichincha. Mt. Pichincha, above Quito along road to the crest. Alt.
3,100 m. Aug 1985. A. Hirtz & W. Flores 2614 (RPSC!). Prov. Sucumbíos. Alt. ca. 3,300 m.
6 July 2017. M. Kolanowska et al. E17/39 (photo!).
The following collections correspond to P. multiflora but the plants have glabrous floral
bracts. In our opinion they should be considered as a separated variety.
Pterichis multiflora var. ecuadorensis Kolan., Szlach. & S. Nowak, var. nov.TYPE:
Ecuador. Prov. Tungurahua. Ambato Cantón. Parque Nacional Llanganates. Cordillera de
los Llanganates. Laguna de Soguillas. Páramo. Perianth mostly yellow with maroon streaks.
Alt. 3,700 m. 27 November 1996. J.L. Clark & J. Fair 3459 (QCNE!). Figs. 5,16 and 17.
Distinguished from typical form of Pterichis multiflora by glabrous floral bracts.
Etymology: In reference to the geographical distribution of the new variety.
Representative specimens: Ecuador. Prov. Azuay. km 50 S of Cuenca on Panamerican
highway. Páramo. Alt. ca. 3,500 m. 6–10 May 1973. Terrestrial in grass paramo.
L.B. Holm-Nielsen et al. 5128 (AAU!). Prov. Carchi. Páramo El Angel. In the pass on road
El Angel-Tulcán. Very humid Espeletia-páramo. Alt. 3,750–3,850 m. 15 May 1973.
L.B. Holm-Nielsen et al. 5450 (AAU!). Prov. Loja. Road Pichig-Fierro Urc, ca. km 10.
Paramo and upper montane forest. Alt. 3,000–3,500 m. 20 Jan 1990. J.E. Madsen 86746
(AAU!). Prov. Napo. Cordillera de los Llanganates, Loma between Río Topo and Río
Verde Grande. 3 km NW of Cerro Hermoso. Alt. 4,000 m. 10 November 1980.
L.B. Holm-Nielsen et al. 28327 (AAU!). Prov. Tungurahua. Ambato Cantón. Parque
Nacional Llanganates. Cordillera de los Llanganates. Laguna de Soguillas. Páramo.
Perianth mostly yellow with maroon streaks. Alt. 3,700 m. 27 November 1996. J.L. Clark &
J. Fair 3459 (QCNE!).
Pterichis parvifolia (Lindl.) Schltr., Bot. Jahrb. Syst. 45: 389. 1911. ≡Acraea parvifolia
Lindl., Pl. Hartw.: 155. 1845. TYPE (Garay, 1978: 186): Ecuador. Loja. Hartweg 50
(lectotype K-L, K-L-Garay’s illustration!).
Pterichis barbifrons (Kraenzl.) Schltr., Repert. Spec. Nov. Regni Veg. Beih. 9: 127.
1921. ≡Prescottia barbifrons Kraenzl., Bot. Jahrb. Syst. 54, Beibl. 117: 19. 1916. TYPE
(Garay, 1978: 186): Peru. A. Weberbauer s.n. (lectotype B; AMES-00103619!—photo,
F-0BN018368!—photo).
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 26/45
Plant 18–45 cm tall. Leaf petiolate; petiole 5.0–6.0 cm long, canaliculated; blade
5.4–7.0 × 1.4–2.0 cm, ligulate to narrowly elliptic, subacute. Scape pubescent, with
4–5 sheaths decreasing in size distally along the scape, pubescent in upper half.
Inflorescence 3.5–7.0 cm long, subdensely 7–10-flowered, rachis pubescent. Flowers
greenish or brownish with yellow lip with brownish veins. Floral bracts 6–12 mm long,
ovate-lanceolate, acute, glabrous. Pedicellate ovary 10.5–21 mm long, pubescent. Dorsal
sepal 8.0–9.0 × 2.3–3 mm, ovate-lanceolate, subacuminate, obtuse, 3-veined, externally
sparsely ciliate. Petals 8–10 × 1.5–2.5 mm, agglutinate to dorsal sepal, unguiculate in
lower fifth, obliquely oblong-lanceolate, subacute, 3-veined, ciliate along margin. Lateral
sepals 8.1–8.5 × 3.6–4.0 mm, obliquely ovate, subacuminate, 3- or 4-veined, externally
Figure 16 Pterichis multiflora var. ecuadorensis Kolan., Szlach. & S. Nowak, var. nov.(A) Floral bract.
(B) Dorsal sepal. (C) Lateral sepal. (D) Petal. (E) Lip. Scale bars = 1 mm. Drawn by S. Nowak from
Madsen 86746 (AAU). Full-size
DOI: 10.7717/peerj.10807/fig-16
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 27/45
sparsely ciliate. Lip 5.1–5.5 × 9.5–10.0 mm, 3-lobed, basal part reniform above truncate
base, with small auricles; middle lobe shorter than lip half length, ovate-triangular, densely
glandular-pubescent; disc ornamented with a knob-like projections along margin,
apical margin ciliate, 7-veined with branching veins. Gynostemium 3.0–4.0 mm long.
Figs. 5 and 18.
Habitat and ecology: Terrestrial in very humid montane forest, in paramo and on steep
mossy exposed slopes without trees at the altitudes of 3,200–3,800 m. Flowering occurs in
June, July and August.
Figure 17 Pterichis multiflora var. ecuadorensis Kolan., Szlach. & S. Nowak, var. nov.(A) Floral bract.
(B) Dorsal sepal. (C) Lateral sepal. (D) Petal. (E) Lip. Scale bars = 1 mm. Drawn by S. Nowak from
Clark & Fair 3459 (QCNE). Full-size
DOI: 10.7717/peerj.10807/fig-17
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 28/45
Notes: This species differs from similar P. habenarioides by the obliquely oblong-lanceolate
(vs. obovate-lanceolate) petals, glabrous (or almost glabrous) sepals (vs. sepals pubescent),
and a presence of a single row of the large papillae along the lip margin (vs. papillae
small, arranged in 2–3rows).Pterichis barbifrons and it was not accepted by most
subsequent researchers as separated species (Schweinfurth, 1958;Garay, 1978).
Unfortunately the type of P. barbifrons was lost and the actual concept of this taxon is
confusing. It was considered as synonym of P. galeata by Schweinfurth (1941) and this
concept was followed by Kolanowska & Szlachetko (2017), however, more detailed
Figure 18 Pterichis parvifolia (Lindl.) Schltr. (A) Dorsal sepal. (B) Lateral sepal. (C) Petal. (D) Lip.
Scale bars = 1 mm. Drawn by S. Nowak from Øllgard & al. 34471 (AAU).
Full-size
DOI: 10.7717/peerj.10807/fig-18
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 29/45
studies of available literature indicated that P. barbifrons has glabrous floral bracts and its
petals are adnate to the dorsal sepal. Hereby this species fits more the morphological
characteristic of P. parvifolia as proposed by Garay (1978).
Representative specimens: Ecuador. Prov. Azuay. Between Cumbe and El Progreso. Alt.
3,200 m. 4 Aug 1975. C. Luer et al. 411 (RPSC!, UGDA-DLSz!—drawing). Prov. Loja.
C.T. Hartweg 50 (K-L-Garay’s illustration!). Prov. Napo. Quijos. Sector de Papallacta.
Sendero hacia las Lagunas de Cojunco y Verde. Alt. 3,350–3,450 m. Bosque muy humedo
subalpino, paramo herbaceo. 25 June 2000. A. Álvarez et al. 2613 (QCNE!). Prov.
Pichincha/Napo. Road Olmedo-Laguna San Marcos, E of the pass. Alt. 3,620–3,800 m.
10–11 July 1980. B. Øllgard et al. 34471 (AAU!).
Pterichis pauciflora Schltr., Repert. Spec. Nov. Regni Veg., Beih. 8: 41. 1921. TYPE (Garay,
1978: 186): Ecuador. Loja. In Andibus orientalibus. Alt. 3,000–3,400 m. F. Lehmann 7111
(B†; lectotype K-000573776!, isolectotype AMES-00083571—fragment).
Plant 14–40 cm tall. Leaf petiolate; petiole 5.0–8.0 cm long; blade 4.0–10.0 cm × 1.2 cm,
linear to oblong-lanceolate, acute. Scape minutely puberulent in the upper part, with
4–5 tubular, pubescent sheaths. Inflorescence 3.0–7.0 cm long, 2–12-flowered, rachis
densely ciliate. Flowers yellowish-orange to yellowish-brown, with brown veins. Floral
bracts up to 8.0 mm long, ovate-lanceolate, glabrous. Pedicellate ovary 8–12 mm long,
densely glandular. Dorsal sepal 5–8×1.3–3 mm, lanceolate-ovate to ovate, obtuse,
3-veined, glabrous or externally very sparsely ciliate. Petals 5.2–7.8 × 1–2.2 mm,
agglutinate to dorsal sepal, obliquely linear-lanceolate, acuminate, l- or 2-veined, margin
sparsely ciliate. Lateral sepals 5–6×2.8–4.5 mm, obliquely ovate, acuminate, 3−veined,
glabrous or externally sparsely ciliate. Lip 4.5–7.5 × 5.5–8.5 mm, basal part reniform
above truncate base; middle lobe indistinctive, triangular, obtuse, glandular-papillose;
disc 7-veined, margin ornamented with large glands. Gynostemium 2.5–3.2 mm long.
Figs. 5 and 19.
Habitat and ecology: Terrestrial in open, humid places in high mountains and in paramo
dominated by Blechnum,Elaphoglossum Schott ex J.Sm. (Lomariopsidaceae), Monticalia
(=Senecio)andicola (Turcz.) C.Jeffrey (Asteraceae), at the altitudes of 3,000–3,910 m.
Flowering occurs in February and April.
Notes: The lip basal part of Pterichis pauciflora is not well-separated from the middle lobe
unlike in the most species of section Acraea. From somewhat similar P. habenarioides and
P. parvifolia it differs also in having usually 1-veined petals. Unlike P. habenaroides
this species has linear-lanceolate petals (Schlechter, 1921). In somewhat similar P. triloba
petals are sparsely ciliate on margin and 3-veined.
Representative specimens: Ecuador. Prov. Azuay. Cuenca Sayaúsi. Parque Nacional
Cajas, cerca del sector San Luis. Alt. 3,910 m. 5 February 2013. D. Minga & A. Verdugo
2530 (HA 8413!). Prov. Loja. In Andibus orientalibus. Alt. 3,000–3,400 m. F. Lehmann
7111 (K-000573776!). Prov. Napo. N side of Cerro Sumaco, upper part of the loma NW od
campsite. Humid páramo in furrow-like quebradas dominated by Blechnum,
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 30/45
Elaphoglossum and Senecio andicola. Alt. 3,750 m. 25 April 1979. L.B. Holm-Nielsen et al.
17247 (AAU!).
Pterichis seleniglossa Schltr., Repert. Spec. Nov. Regni Veg. Beih. 8: 42. 1921. TYPE:
Ecuador. Pichincha. Ad rupes montium. Alt. 3,300 m. L. Sodiro s.n. (B?).
Plant 16–35 cm tall. Leaf petiolate; petiole 2.0–10.0 cm long; blade 5.0–16.0 ×
1.5–2.3 cm, linear to oblong-lanceolate, acute. Scape, minutely puberulent in the upper
part, with 3–6 tubular, glandular sheaths. Inflorescence 5.0–7.5 cm long, sublaxly to
Figure 19 Pterichis pauciflora Schltr. (A) Dorsal sepal. (B) Lateral sepal. (C) Petal. (D) Lip. Scale
bars = 1 mm. Drawn by S. Nowak from Holm-Nielsen al. 17247 (AAU).
Full-size
DOI: 10.7717/peerj.10807/fig-19
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 31/45
subdensely 8-13-flowered; rachis densely ciliate. Flowers with dark green or brown sepals
and petals and brown or yellow lip marked with red-brown or yellow. Floral bracts
7.0–8.0 mm long, ovate-lanceolate, glandular-ciliate. Pedicellate ovary 8.0–12.0 mm long,
densely glandular-ciliate. Dorsal sepal 6.0–9.0 × 1.6–3.2 mm, ovate-lanceolate to ovate,
obtuse, 3- or 5-veined, externally ciliate. Petals 6.5–9.0 × 1.7–2.2 mm, agglutinate to dorsal
sepal, obliquely linear-lanceolate or linear, acuminate, obtuse, l-veined, glabrous. Lateral
sepals 6.0–8.0 × 2.1–3.5 mm, obliquely ovate-lanceolate to ovate, obtuse, 3-veined,
externally ciliate. Lip 3.2–6.8 × 7.5–9.0 mm, basal part lunate above truncate base; middle
lobe shorter than lip half length, elliptic, obtuse, papillate; disc with numerous small glands
along the margin, 7- or 9-veined, veins apically branching. Gynostemium 2.5–3.0 mm
long. Figs. 5 and 20.
Habitat and ecology: Terrestrial in high-montane forest and along roadside in grass
paramo at the altitudes of 3,050–3,300 m. Flowering occurs in March, July and August.
Figure 20 Pterichis seleniglossa Schltr. (A) Floral bract. (B) Dorsal sepal. (C) Lateral sepal. (D) Petal.
(E) Lip. Scale bars = 1 mm. Drawn by S. Nowak from Dodson & al. 16396 (QCNE).
Full-size
DOI: 10.7717/peerj.10807/fig-20
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 32/45
Notes: Schweinfurth (1958) and Garay (1978) considered this species synonymous to
Pterichis triloba. Based on our study the two species differs in petal venation. Petals of
P. seleniglossa are 1-veined (vs. 3-veined in P. triloba). Pterichis seleniglossa resembles
Bolivian P. mandonii Rolfe but in the latter species the lip middle lobe is much shorter.
Representative specimens: Ecuador. Prov. Bolivar. Guaranda-Caluma-Catarama, km 7.7.
Alt. 3,050 m. 5 Jul 1991. C.H. Dodson, N. Williams & M. Whitten 18767 (RPSC!,
UGDA-DLSz!—drawing). Prov. Pichincha. Yanacocha Reserve. 12 August 2018.
M. Kolanowska, S. Nowak & A. Hirtz E18/14 (photo!), Reserva Gebotánica del Pululahua.
Alt. 3,100 m. 1 March 2001. C. Cerón & E. Freire 43973 (QAP 38931!); Ad rupes montium.
Alt. 3,300 m. L. Sodiro s.n.(Schlechter, 1921). Prov. Imbabura. Road from Otavalo to
Selva Alegre via Lago Cuicocha entering from Panamerican highway between Otavalo and
Ibarra. Alt. 3,100 m. 20 March 1986. C.H. Dodson et al. 16396 (QCNE!).
Pterichis triloba (Lindl.) Schltr., Bot. Jahrb. Syst. 45: 389. 1911. hAcraea triloba Lindl.,
Ann. Mag. Nat. Hist. 15: 386. 1845. TYPE (Garay, 1978: 187): Ecuador. Prov. Pichincha.
Hda. de Pinantura, 35 km SE of Quito. C.T. Hartweg s.n. (lectotype K-L, K-L-Garay’s
illustration!, isolectotype AMES-00082267!—fragment).
Plants 27–34 cm tall. Leaf sessile, (6.0)16.0–24.0 cm long, narrowly lanceolate,
acuminate. Scape ciliate in the upper half, enclosed with 4–5 tubular sheaths which are
gradually shorter towards the scape. Inflorescence 4.0–8.0 cm long, sublaxly to subdensely
7–15-flowered, rachis ciliate. Flowers with greenish-brown sepals and petals and basally
red-brown lip with yellow apex. Floral bracts 7.0–11.0 mm long, ovate or ovate-lanceolate,
ciliate. Pedicellate ovary 10.0–14.0 mm long, pubescent. Dorsal sepal 7.0–10.0 × 1.8–3.1
mm, elliptic-lanceolate, obtuse, 3-veined, externally ciliate. Petals 7.0–10.0 × 1.0–3.0 mm,
agglutinate to dorsal sepal, oblong-elliptic to oblong-lanceolate, obtuse, 3-veined,
glabrous. Lateral sepals 6.0–9.3 × 2.0–3.2 mm, obliquely elliptic-ovate, subacute, 3-veined,
externally ciliate. Lip 5.1–6.7 × 7.5–9.0 mm, distinctly 3-lobed, basal part reniform-flabellate
above truncate base; middle lobe ca 1/3 of the lip length, elliptic, apiculate; disc glandular
with numerous glands along margin. Gynostemium 2.2–4.0 mm long. Figs. 5 and 21.
Habitat and ecology: Terrestrial on steep slopes in high mountains and in mossy scrub
forest and open grasslands at the altitudes of 3,100–3,600 m. Flowering occurs in March,
April, May and July.
Notes: This species resembles somewhat P. tunguraguona in which the truncate base of the
lip is very short (less than 1/3 of the lip lamina vs. 1/3 of the lip lamina) and there are no
glands or projections along the margin of the basal lip part.
Representative specimens: Ecuador. Prov. Imbabura. Laguna Culcocha, crater lake 30 km
W of Ibarra. 3,100 m. 24 May 1973. Terrestrial on a steep rocky slope. L.B. Holm-Nielsen
et al. 6370 (AAU!). Prov. Pichincha. 30 March 1930. R. Benoiss 2315 (Q 18090!); Quito-Santo
Domingo via Chiriboga, km 20. Alt. 3,400 m. 15 May 1981. Terrestrial in paramo.
C.H. Dodson et al. 10837 (Q!, QCA!, RPSC!); Antisana, camino entre Inga Montserrat y
Vaquero Pasana. Alt. 3,400–3,600 m. 13 April 1992. A. Fieire-Fierro et al. 2143 (NY!, QCA!);
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 33/45
Mt. Mojanda, at the TV antenna on top of Mt Mojanda entering from km 10 on the
Otavalo-Selva Alegre road. Alt. 3,100–3,400 m. 31 July 1985. C.H. Dodson & A. Embree
16121 (RPSC!, UGDA-DLSz!—drawing); Hda. de Pinantura, 35 km SE of Quito.
C.T. Hartweg s.n.(K-L–Garay’s illustration!, AMES!); Volcan Corazón. Alt. 3330. 4 July
2017. M. Kolanowska et al. E17/10 (photo!).
Pterichis tunguraguona Rchb. f. ex Szlach. & Kolan., Ann. Bot. Fenn. 51(5): 331. 2014.
TYPE: Ecuador. W. Jameson s.n. (holotype W 878!).
Plant 16–40 cm long. Leaf petiolate; petiole 2.8–7.0 cm long; blade 5.0–14.0 ×
1.3–3.0 cm, narrowly elliptic, obtuse to acute. Scape glabrous, enclosed in 3–5 tubular
Figure 21 Pterichis triloba (Lindl.) Schltr. (A) Dorsal sepal. (B) Lateral sepal. (C) Petal. (D) Lip. Scale
bars = 1 mm. Drawn by S. Nowak from Benoiss 2315 (Q). Full-size
DOI: 10.7717/peerj.10807/fig-21
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 34/45
sheaths, the uppermost sheaths often ciliate. Inflorescence 4.5–17.0 cm long, subdensely to
sublaxly 7–15-flowered. Flowers white-yellow or olive-green with weak brown-green stipes
sepals, yellowish petals which are basally red-brown tinted, and brown-yellow or greenish
yellow lip with dark brown stripes, apex pale yellow. Floral bracts 7.0–12.0 mm long,
lanceolate to elliptic, sparsely ciliate. Pedicellate ovary 7.0–13.0 mm long, glandular-ciliate.
Dorsal sepal 6.1–8.8 × 1.9–2.8 mm, oblong-lanceolate to ovate, obtuse, 3- or 5-veined,
externally papillate. Petals 7.0–9.0 × 2–4 mm, agglutinate to dorsal sepal, narrowly ovate
above narrow claw, obtuse, 3-veined, glabrous. Lateral sepals 7.0–9.5 × 3.0–3.5 mm,
obliquely ovate-lanceolate, obtuse, margins incurved, 3-veined, externally papillate.
Lip 5.8–6.5 × 8.0–9.5 mm, basal part reniform-lunate above truncate base, with small
auricles; middle lobe ca 1/3 of the lip length, ligulate, papillate; disc papillate, with
numerous small glands along the margins, 7- or 9-veined, veins apically branching.
Gynostemium 2.2–2.5 mm long. Figs. 5 and 22.
Habitat and ecology: Terrestrial in paramo, in montane forest and on the trails at the
altitudes of 2,900–3,660 m. Flowering occurs in January, May and September.
Notes: This is one of four Ecuadorian representatives of sect. Acraea with glabrous petals.
It can be distinguished from Pterichis seleniglossa and P. triloba by relatively short
truncate basal part of the lip (less than 1/3 of the lip lamina). Pterichis tunguraguona is
resembles P. parviflora but the petals of P. tunguraguona are shortly unguiculate at base
(vs. non-unguiculate), narrowly ovate above (vs. narrowly-oblong), distinctly wider
than sepals (vs. narrower than sepals). Moreover, its lip is transversely elliptic, semilunate
at the base (vs. lip basal part semiorbicular) and papillate along apical margins and on the
apex, and the lip apical part is ligulate-lanceolate (vs. triangular). Distal lip margins are
papillate and densely ornamented by numerous, swollen outgrowths, which are missing in
P. tunguraguona.
Representative specimens: Ecuador. Prov. Azuay. km 91 on Panamaerican Highway N of
Loja. Dry scrub. Alt. 2,900 m. 5 May 1973. L.B. Holm-Nielsen et al. 5108 (AAU!). Prov.
Imbabura. Vía Joya-Laguna de Mojanda Cajas. Al sur de la población de Otavalo. Alt.
3,109–3,659 m. 26 January 1980. J. Jaramillo & F. Coello 2042 (AAU!, QCA!); Above
Piñán, slopes of Volcan de Cotacachi. Alt. 10,450 ft. 3 September 1944. W. B. Drew E-91
(QCNE!). Sine loc.W. Jameson s.n. (W!).
Pterichis hirtziana Kolan., Szlach. & S. Nowak, sp. nov.TYPE: Ecuador. Villonaco. Ca.
15 km W of Loja. Alt. 2,500–2,900 m. 24 April 1987. H. van der Werff & W. Palacios 8956
(holotype: QCNE 34289!).
Species similar to Pterichis habenarioides distinguished by lunate basal lip part and
obliquely oblong-elliptic petals.
Plants about 25 cm tall. Leaf not observed in herbarium material. Scape glandular in the
upper part, enclosed in 6 tubular sheaths. Inflorescence 6.0 cm long, 12-flowered, rachis
densely ciliate. Flowers greenish-red. Floral bract 8.2 mm long, very sparsely ciliate.
Pedicellate ovary 11.0 mm long, ovate to broadly ovate, ciliate. Dorsal sepal 8.3 × 2.2 mm,
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 35/45
oblong-ovate, rounded at the apex, 5-veined, glabrous. Petals 8.1 × 1.8 mm, agglutinate
to dorsal sepal, obliquely oblong-elliptic, obtuse, 3-veined, sparsely ciliate. Lateral sepals
7.1 × 3.0 mm, broadly ovate, acuminate, subobtuse, 3-veined, glabrous. Lip 5.5 × 8.8 mm,
basal part lunate above truncate base; middle lobe ca 1/3 of lip length, oblong-ovate,
obtuse, densely papillate; disc sparsely papillate, ornamented with numerous small
glands along margins, 7-veined, veins apically branching. Gynostemium 3.0 mm long.
Figs. 5 and 23.
Etymology: Dedicated to Alexander Hirtz, an eminent Ecuadorian orchidologist.
Habitat and ecology: Terrestrial in cloud forest changing into shrubby paramo at the
altitudes of 2,500–2,900 m. Flowering occurs in April.
Notes: This species resembles Pterichis habenarioides in having sparsely ciliate floral bracts,
glabrous sepals and sparsely ciliate petals, but it is easily distinguished from the latter by
Figure 22 Pterichis tunguraguona Rchb. f. ex Szlach. & Kolan. (A) Lateral sepal. (B) Dorsal sepal.
(C) Petal. (D) Lip, front view. (E) Lip, lateral view. Scale bars = 1 mm. Drawn by A. Król from Jameson s.
n. (holotype W). Full-size
DOI: 10.7717/peerj.10807/fig-22
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 36/45
having lunate lip basal part (vs. reniform), and obliquely oblong-elliptic petals
(vs. lanceolate).
Representative specimen: Ecuador. Prov. Loja. Villonaco. Ca. 15 km W of Loja. Alt.
2,500–2,900 m. 24 April 1987. H. van der Werff & W. Palacios 8956 (QCNE! 34289).
Pterichis dodsoniana Kolan., Szlach. & S. Nowak, sp. nov.TYPE: Ecuador. Prov.
Azuay. Km 85 on Panamerican highway N of Loja. Alt. 2,850–2,950 m. 3 May 1973.
L.B. Holm-Nielsen et al. 4767 (holotype AAU!; isotype AAU!).
Species similar to Pterichis habenarioides but with almost linear petals which are longer
than dorsal sepal and prominent lip middle lobe.
Plants 24–29 cm tall. Leaf petiolate; petiole 7.5 cm long; blade 6.8 cm long, narrowly
lanceolate. Scape ciliate above basal sheath, with 3–4 tubular sheaths, the upper one
sparsely ciliate. Inflorescence 4.0–10.0 cm long, 4–12-flowered, rachis densely ciliate.
Figure 23 Pterichis hirtziana Kolan., Szlach. & S. Nowak, sp. nov.(A) Floral bract. (B) Dorsal sepal.
(C) Lateral sepal. (D) Petal. (E) Lip. Scale bars = 1 mm. Drawn by S. Nowak from van der Werff &
Palacios 8956 (QCNE). Full-size
DOI: 10.7717/peerj.10807/fig-23
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 37/45
Flowers greenish-yellow with green veins, petals yellow with brown veins. Floral bract
6.6 mm long, glabrous. Pedicellate ovary 10.0 mm long, ovate to broadly ovate, pubescent.
Dorsal sepal 7.8 × 2.8 mm, narrowly ovate, obtuse, 5-veined, glabrous. Petals 8.2 × 1.2 mm,
agglutinate to dorsal sepal, linear, widened in the middle, obtuse, 3-veined, sparsely
ciliate. Lateral sepals 6.3 × 3.1 mm, ovate, acuminate, subacute, 3-veined, externally ciliate
only near the base. Lip 7.0 × 8.7 mm, basal part reniform above truncate base, with small
auricles; middle lobe ca 1/3 of the lip length, ligulate-ovate, apiculate, obtuse, densely
papillate; disc sparsely papillate, ornamented with numerous glands along margins,
7-veined. Gynostemium 3.2 mm long. Figs. 5 and 24.
Etymology: Dedicated to Dr. Calaway Dodson (1928–2020), eminent orchid taxonomist,
who described numerous orchid species from Ecuador. He was the author of partial
treatments of Orchidaceae for the Flora of Ecuador project of Gothenburg University
(Sweden) and the Universidad Catolica of Ecuador.
Figure 24 Pterichis dodsoniana Kolan., Szlach. & S. Nowak, sp. nov.(A) Floral bract. (B) Dorsal sepal.
(C) Lateral sepal. (D) Petal. (E) Lip. Scale bars = 1 mm. Drawn by S. Nowak from Holm-Nielsen & al. 4767
(AAU). Full-size
DOI: 10.7717/peerj.10807/fig-24
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 38/45
Habitat and ecology: Terrestrial in dry, low scrub vegetation at the altitudes of
2,850–2,950 m. Flowering in May.
Notes: This species resembles Pterichis habenarioides in having glabrous floral bracts and
sepals and in sparsely ciliate petals, but it is easily distinguished from the latter by
having linear petals slightly widened in the middle (vs. lanceolate) which are longer than
dorsal sepal, and prominent lip middle lobe constituting more than 1/3 of the total lip
length (vs. less than 1/4 of the total lip length).
As holotype serves specimen in the middle part of herbarium sheet, two other plants are
therefore isotypes.
Representative specimen: Ecuador. Prov. Azuay. km 85 on Panamerican highway N of Loja.
Alt. 2,850–2,950 m. 3 May 1973. L.B. Holm-Nielsen et al. 4767 (AAU!).
Incertæ sedis
Material incomplete or too damaged for identification
Ecuador. Prov. Azuay. Cantón Sigsig. Parroquia Jima, parte alta de Moya, sector
Zhurugiña. Alt. 3,400 m. L. Suin et al. 1041 (HA!). Prov. Carchi. Cantón Mantufar,
San Gabriel, Loma Bretaña, entre Río Minas y la quebrada Central. Alt. 3,100–2,500 m.
18 August 2010. C. Cerón et al. 68580 (QAP!). Prov. Cotopaxi. Cantón Sichchos, sector
Monte Cimaurcu. Alt. 4,030 m. 2 June 2012. C. Cerón & D. Jácome 71627 (QAP!).
Prov. El Oro. Chilla Cantón. Cerro de Chilla. Vegetación de paramo, arbustiva y herbacea.
Alt. 3,595 m. 3 May 1997. H. Vargas & C. Conaday 1393 (QCNE!). Prov. Imbabura.
Alturas de Cayachupa, ca 5 km NW of the village Piñan. Alt. 3,200 m. 16 June 1980
C. R. Sperling & R. Bleiweiss 5105 (QCA!). Prov. Loja. Parque Nacional Podocarpus. Cerro
Toledo. 1 December 1988. J. E. Madsen & al 75700 (AAU!). Prov. Napo. Parque Nacional
Llanganates. Vía Salcedo-Tena, sector Siete vueltas. Alt. 3,600–3,850 m. 26 February 2015.
A.J. Pérez et al. 8317 (QCA!). Prov. Pichincha, cantón Quito. Reserva Geobotánica del
Pululahua, sector Lulumbamba. Alt. 2,700 m. 7 June 2008. C. Cerón & C. Reyes 62369
(QAP!), Cantón Quito. Reserva Geobotánica del Pululahua, Cerro Padre Rumi. Alt. 2,825 m.
2 June 2007. C. Cerón & C. Reyes 59179 (QAP!). Prov. Zamora-Chinchipe. Cordillera de
Sabanilla, cerca de la carretera Jimbura-Zumba. Alt. 3,300–3,500 m. 22 October 1996.
RBu & SL 1221 (QCA!). Without specific locality. Ad rupes Cotursi –Pifi. Alt. 3,000 m.
A. Mille 31 (QPLS!).
DISCUSSION
Globally orchids are among the most threatened plants due to increase of anthropogenic
pressures, high level of endemism and complex relationships with mycorrhizal fungi
and pollinators (Wraith, Norman & Pickering, 2020). As suggested by Pillon & Chase
(2007), taxonomic recognition of orchids is crucial for planning any conservation actions
to protect these plants and species inventories, especially in the most biodiverse tropical
areas, are required establish adequate conservation projects (May, 1988;Dubois, 2003).
The systematic surveys are particularly important considering the crisis of biodiversity
(Ramade, 1999) which can lead to extinction of numerous rare organisms. Unfortunately,
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 39/45
also the taxonomic research are in crisis (Tillier, 2000;Dubois, 2003). The lack of clear
characteristic of particular species results in numerous misidentifications of known taxa
and false local species checklists (Dubois, 2003).
According to our findings only floral bracts and perianth segments are diagnostic
characters that allow to identify Pterichis species, however, the importance of several
characters requires further studies—the genetic analyses could help resolve some
classification doubts. For example we found that P. acuminata is characterized by a
significant variation and we are not able to clearly state that small deviations from the
typical morphology of the floral segments are enough to recognize some populations as
separated species (see notes in P. acuminata). The ornamentation of floral bracts seems
to be consistent within the same species and the bracts are either ciliate or glabrous.
Rarely intermediate form of floral bract, with just few ciliae, was observed. The similar
scheme was recorded for petals and sepals which shows the same pattern of ornamentation
within the species (but see notes in P. acuminata). The lip seems to be the most
conservative character—its form, relative size and distribution of papillae or knobs is
consistent in populations of the same species. The venation of tepals is the most disputable
diagnostic character and it requires further analyses. While usually petals of Pterichis
representatives are 1- or 3-veined and sepals are 3- or 5-veined, some populations with
2- or 4-veined petals were found during our research. Our new species are described based
on several (at least three) differences between novelties and the most similar taxa.
In this article the new species are described based on a single or just several plants of the
same collection, however, it is not an unusual situation. Preferably any new taxon should
be formally named after the examination of numerous randomly collected specimens
from various populations (Dubois, 2010). Nevertheless, a considerable number of species,
not only plants, are described based on a single specimen (Captain et al., 2019;Wells,
Johanson & Dostine, 2019). The disadvantage of using a few specimens for the description
of new taxa, particularly of new species, is that the intraspecific variation cannot be
evaluated. As calculated by Raven et al. (2020) an average of 749 new vascular plant species
has been described annually from Latin America for the past 25 years and this rate
did not decrease with time. According to some estimates (Bebber et al., 2010) about
70,000 species of flowering plants are waiting to be discovered and named. Around half
of the anticipated missing species presumably have already been collected and these
are deposited in herbaria awaiting identification (Bebber et al., 2010). On the other
hand, Fontaine, Perrard & Bouchet (2012) calculated that an average time between the
first collection of a specimen of a new species to its formal description is 21 years.
Considering the ongoing habitat loss it is not therefore surprising that numerous novelties
found in herbarium are difficult to locate in its natural habitat and for that reason the number
of samples used in new species description is often limited. According International Plant
Name Index (IPNI) only in 2019 more than 150 new Orchidaceae species were described.
Novelties were found mainly in tropical in subtropical regions and numerous new names, for
example Dichaea amazonica Pupulin, Epidendrum brevicallosum Hágsater & E. Santiago,
were published based on single specimen. By examination of numerous Pterichis from various
geographical regions (Kolanowska & Olędrzy
nska, 2015;Kolanowska & Szlachetko, 2017)
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 40/45
we were able to recognize the diagnostic characters of this genus representatives and only
these taxonomically important traits were used to describe new species.
The last study which included the recognition of Pterichis diversity in Ecuador included
only six species (Garay, 1978). In contrast to Garay’s (1978) concept which did not
recognize P. seleniglossa as separated species, in our opinion this taxon clearly differs from
all other genus representatives and can be distinguished from similar P. triloba by 1-veined
petals (vs. 3-veined). In our study we did not find any population of P. triloba or
P. seleniglossa with intermediate, 2-veined petals hereby in our opinion this character is
sufficient for species separateness.
CONCLUSIONS
Here we presented the synopsis of Ecuadorian representatives of Pterichis. In our
opinion species characteristics and identification keys will be useful for local scientists
working on local floras inventories and for launching more sophisticated nature
management programs.
We confirmed the occurrence of 17 Pterichis species in Ecuador. That number includes
four new species described in this article. Seven genus representatives are endemic and
were not so far found outside the country—P. ansaloniana, P. dodsoniana, P. elliptica,
P hirtziana, P. madsenii, P. meirax, and P. tunguraguona. National Pterichis occur in just
three ecoregions (Eastern Cordillera real montane forests, Northern Andean páramo,
Northwestern Andean montane forests) and they are growing usually as terrestrial herbs
between 2,300 and 4,110 m a.s.l.
ACKNOWLEDGEMENTS
We are grateful to the Curators of the cited herbaria and collaborators of these institutions
for access to their specimens or for sending specimens on loan. We would like to thank
Leonardo Guimarães and two anonymous Reviewers for their valuable comments on
the manuscript.
ADDITIONAL INFORMATION AND DECLARATIONS
Funding
This work was supported by the Ministry of Education, Youth and Sports of Czech
Republic within the CzeCOS program, grant number LM2018123 and by the National
Science Centre, Poland (PRELUDIUM 2016/21/N/NZ8/01601). The funders had no role
in study design, data collection and analysis, decision to publish, or preparation of the
manuscript.
Grant Disclosures
The following grant information was disclosed by the authors:
Ministry of Education, Youth and Sports of Czech Republic: LM2018123.
National Science Centre, Poland: PRELUDIUM 2016/21/N/NZ8/01601.
Kolanowska et al. (2021), PeerJ, DOI 10.7717/peerj.10807 41/45
Competing Interests
Marta Kolanowska is an Academic Editor for PeerJ.
Author Contributions
Marta Kolanowska conceived and designed the experiments, performed the
experiments, analyzed the data, prepared figures and/or tables, authored or reviewed
drafts of the paper, and approved the final draft.
Dariusz L. Szlachetko conceived and designed the experiments, performed the
experiments, analyzed the data, authored or reviewed drafts of the paper, and approved
the final draft.
Sławomir Nowak conceived and designed the experiments, performed the experiments,
analyzed the data, prepared figures and/or tables, authored or reviewed drafts of the
paper, and approved the final draft.
Data Availability
The following information was supplied regarding data availability:
This is taxonomic revision; all data (e.g., list of herbairum specimens examined) are
presented in the article in the sections entitled “Representative Specimens”under
characteristic of each taxon.
New species are deposited:
Pterichis ansaloniana—C. Ulloa et al. 1361 (HA 5333)
Pterichis madsenii—J.E. Madsen & L. Ellemann 75293 (AAU s.n.).
Pterichis hirtziana—H. van der Werff & W. Palacios 8956 (QCNE 34289).
Pterichis dodsoniana—L.B. Holm-Nielsen et al. 4767 (AAU s.n.).
New Species Registration
The following information was supplied regarding the registration of a newly described
species:
Pterichis ansaloniana Kolan., Szlach. & S. Nowak, sp. nov.: 77214449-1
Pterichis madsenii Kolan., Szlach. & S. Nowak, sp. nov.: 77214450-1
Pterichis hirtziana Kolan., Szlach. & S. Nowak, sp. nov.: 77214451-1
Pterichis dodsoniana Kolan., Szlach. & S. Nowak, sp. nov.: 77214452-1
Pterichis elliptica var. ecuadorensis Kolan., Szlach. & S. Nowak, var. nov.: 77214453-1
Pterichis multiflora var. ecuadorensis Kolan., Szlach. & S. Nowak, var. nov.: 77214454-1
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