Plant Syst Evol
DOI 10.1007/s00606-014-1054-4
ORIGINAL ARTICLE
Eight new species of Gomphichis (Orchidaceae, Spiranthoideae)
from Colombia
Sławomir Nowak • Dariusz L. Szlachetko •
Joanna Mytnik-Ejsmont • Antoine M. Cleef
Received: 3 June 2013 / Accepted: 21 March 2014
The Author(s) 2014. This article is published with open access at Springerlink.com
Abstract Eight new species of the genus Gomphichis
from Colombia are described. Each species is illustrated,
and detailed habitat and distribution data are provided. A
distribution map of the new species is presented. A
dichotomous key for determination of the Gomphichis
species in northern South America is provided. Conservation status assessments are provided for each species;
current International Union for Conservation of Nature
(IUCN) Red List categories and criteria are listed. A brief
discussion of spiranthoid orchids taxonomy, conservation
status of species, endemism in the Andes and paramo are
presented.
Keywords Andes IUCN categories Orchidaceae
Gomphichis Taxonomy Paramo
Introduction
Field expeditions and botanical explorations of Colombian
flora started in the second half of the eighteenth century;
however, Colombia is still one the South American countries lacking a complete orchid flora and one of the most
megadiverse countries considering its flora (Forero 1988;
Mittermeier et al. 2004). It is estimated that Colombia
S. Nowak (&) D. L. Szlachetko J. Mytnik-Ejsmont
Department of Plant Taxonomy and Nature Conservation,
Faculty of Biology, University of Gdańsk, ul. Wita Stwosza 59,
80-308 Gdańsk, Poland
e-mail: slawomir.nowak@ug.edu.pl
A. M. Cleef
Institute for Biodiversity and Ecosystem Dynamics,
Faculty of Science, University of Amsterdam, Science Park 904,
1098 XH Amsterdam, The Netherlands
harbours about 10 % of the world’s biodiversity (Miani and
Fajardo 2001), including almost 25,000 vascular plant
species, which gives the country the second place considering the diversity of flora (Bernal et al. 2007). Approximately, nearly 30 % of native representatives of
Colombian plants are considered to be endemic (Davis
et al. 1997; Porup et al. 2009). Orchids, occurring in almost
every habitat, are as much as 10 % of the vascular plants of
this country (Kress 1986).
The main factor driving Colombia’s biodiversity richness is the specific geographic position in northwestern
South America. Colombia is the only country in South
America bordering Panama; thus its territory is a meeting
place of representatives of the fauna and flora of both
American continents. Furthermore, this region has a great
climatic and topographic complexity resulting from the
presence of the Andean mountain ranges crossing the
country and influence of both the Caribbean Sea and
the Pacific Ocean. A landscape variety of Colombia and
significant differences in local habitat conditions associated
with substantial differences in elevation, temperature,
precipitation and sun exposure allow the occurrence of
virtually every ecosystem of the planet known (Miani and
Fajardo 2001).
Gomphichis Lindl. is one of the orchid genera occurring
in Colombia. It is a genus including about 30 species
reported from Costa Rica, Brazil, Guyana and western
South America from Venezuela to Bolivia. In the most
recent work, (Ortiz Valdivieso and Uribe Vélez 2007)
noted 13 species for Colombia, but in our opinion this
number is underestimated, as evidenced by this and previous work (Szlachetko et al. 2013). The representatives of
Gomphichis inhabit mountain regions mainly between
1,700 and 3,600 m a.s.l. They occur on damp grassy slopes,
cliffs, in thickets, woodland margins, along roadsides,
123
S. Nowak et al.
upper montane forest and even in open Andean páramo
(Pridgeon et al. 2003).
Dressler (1990) placed Gomphichis within the subtribe
Prescottiinae Dressler with few mainly high Andean genera
(Aa Rchb.f., Altensteinia Kunth, Myrosmodes Rchb.f.,
Porphyrostachys Rchb.f., Prescottia Lindl., and Stenoptera
C.Presl) distinguishing them from Cranichidinae Lindl. by
having, Spiranthes-type velamen, laminar rostellum, soft
pollinia, and lack of humulus (Álvarez and Cameron 2001;
Álvarez-Molina and Cameron 2009; Dressler 1990, 1993;
Szlachetko and Rutkowski 2000). Szlachetko (1995)
accepted the conception of Prescottiinae proposed by
Dressler (1990), but suggested adding Pseudocranichis
Garay to this subtribe and to rank all Prescottiinae under
the tribe Spirantheae Endl.
The subtribe Prescottiinae is a subject of an ongoing
taxonomic work conducted by our team for years (MytnikEjsmont et al. 2012; Szlachetko et al. 2013; Szlachetko and
Nowak 2013). The collected specimens are identified based
on the comparison with the type material and original
descriptions. The results of our recent herbarium studies
(2009–2012) revealed the discovery of eight new species of
Gomphichis in Colombia and they are presented in this
paper.
Materials and methods
The examined material was prepared by using a stereomicroscope with standard classical taxonomy methods. The
morphological descriptions and comparisons were based on
the study of Gomphichis specimens from AMES, B, BM,
COL, CUVC, K, MO, P, VALLE, W and those collected
during the field expeditions by Szlachetko (2005–2013), and
Kolanowska (2009–2011). Standard procedure of preparing
the herbarium material to facilitate stereomicroscopic observation was applied. The following vegetative characters of
individual plants were analysed: stem (height, shape, presence
of glandular hairs), leaves (number, size, shape), sheaths
(number, shape, size), inflorescence (size, density), floral
bracts (size, shape, presence of glandular hairs), flowers,
taken from the middle part of the inflorescence (size of pedicel and ovary, presence of mentum, size and shape of lateral
sepals, dorsal sepal, petals, and lip), as well as gynostemium
(height and shape of column, presence of column foot, pattern
of hair cover of column part). Particular parts of the flower
were rehydrated, dissected, measured and drawn under a
stereomicroscope. The results were then analyzed and compared with the type material, diagnoses and original illustrations. The database of the drawings and photographs of all
studied specimens are available in the first author’s archives.
A key for determination of the Gomphichis species in
northern South America is dichotomous. Acronyms of
123
herbaria followed Holmgren and Holmgren (1998). To draw
the distribution map and prepare line drawings, CorelDRAW
Graphics Suite 12 was used. The IUCN categories were
assigned according to IUCN (2011) guidelines.
Taxonomic treatment
A key to the Gomphichis species in northern South
America
1. Lip sessile or clawed, in the latter case claw very short
and wide………………………………………………..2
1* Lip clawed, claw narrow………………………….16
2. Petals as wide as or wider than dorsal sepal……….3
2* Petals distinctly narrower than dorsal sepal……….9
3. Petals elliptic or oblong–elliptic with broad base,
sessile…………………………………………………..4
3* Petals elliptic–obovate or obovate, attenuate towards
base, shortly petiolate…………………………………..5
4. Lip with prominent furrow along midnerve, with
two fleshy and thick keels along its margins,
gynostemium pubescent dorsally, ciliate ventrally…………………………G. pseudogoodyeroides
4* Lip with massive cushion-like thickening in the
apical half, which is sulcate in front, gynostemium
ciliate on both surfaces…………………G. salamancae
5. Petals densely pubescent on the outer surface………6
5* Petals glabrous or sparsely pubescent on the outer
surface………………………………………………….8
6. Lip widest near the middle, attenuate gradually
towards triangular apex, gynostemium densely pubescent on both surfaces………………………………..…7
6* Lip widest at the rounded apex, attenuate towards the
base, shortly mucronate, gynostemium ciliate at the base
on the ventral surface………………….…G. epiphytica
7* Lip with massive cushion-like inconspicuously
sulcate thickening in apical half……………G. altissima
7* Lip with disc thickened along midvein, margins
membranaceous……………………………….…G. alba
8. Leaves linear–lanceolate, lip widest at base, broadly
obovate in outline, apical half with two cushion-like
thickening separated by furrow, petals pubescent along
margins……………………………...…G. carlos-parrae
8* Leaves obliquely elliptic–lanceolate, lip widest near
the middle, pentagonal in outline, with thickened central
and apical part, petals ciliate along margins…G. viscosa
9. Lip with large or small, but prominent subglobose or
oblong basal thickenings…………………………...…10
9* Lip without any basal thickenings…………………15
10. Basal thickenings massive, very large, 1/3 of the
whole lip length……………………..…G. magnicallosa
10* Lip basal thickenings much smaller…………..…11
Eight new species of Gomphichis
11. Lip narrowly obovate…………………………..…12
11* Lip not as above…………………………………13
12. Lip widest near the middle, apical part densely
pubescent on the inner surface, petals ciliate on both
margins, 1-nerved………………………...…G. renziana
12* Lip widest at the base, apical part papillate,
petals pubescent on both margins, obscurely 3-nerved……………………………………………G. valida
13. Petals long-pubescent along inner margin and ciliate
along outer margins, 1-nerved………..…G. fernandezii
13* Petals ciliate on the outer margins, 3-nerved, rarely
obscurely 3-nerved……………………………………14
14. Lip as long as wide or wider……………G. traceyae
14* Lip distinctly longer than wide……G. goodyeroides
15. Lip oblong–ligulate in outline, densely pubescent
on margins, cochleate and slightly thickened at
apex…………………………………...…G. schneideri
15*Lip not as above………………………G. jaramilloi
16. Lip wider than long………………………………17
16* Lip longer than wide…………………………….18
17. Lip lateral lobes obliquely orbicular, petals ciliate
along margins…………………………...…G. longifolia
17* Lip lateral lobes obliquely triangular, petals ciliate
along both margins near the middle, base and apex
glabrous……………………………….…G. monstruosa
18. Petals wider than sepals…………………………..19
18* Petals narrower than sepals………………………22
19. Petals prominently clawed………………………..20
19* Petals sessile, with broad base……………………21
20. Petals and sepals subacute, sepals 1-nerved……………………………………………G. lozanoi
20* Petals and sepals obtuse, obscurely 3-nerved………………………………….…G. brachystachys
21. Petals suborbicular–elliptic, lip lateral lobes obliquely
rhombic–elliptic, truncate at apex………G. costaricensis
21* Petals elliptic–lanceolate, Lip lateral lobes elliptic,
rounded at apex…………………………G. hetaerioides
22. Petals widest near the middle, attenuate towards
acute apex and towards base…………………………23
22* Petals more or less oblong–elliptic to linear–ligulate,
rounded at the apex……………………………………26
23. Lip oblong–ligulate, almost as wide at the base as
near the apex…………………………..…G. crassilabia
23*Lip more or less pentagonal in outline, distinctly
wider at base than at the apex…………………………24
24. Lip lateral lobes triangular, subacute……G. scaposa
24* Lip lateral lobes rounded…………………………25
25. Petals pubescent on both margins, ciliate on the outer
surface……………………………………G. lancipetala
25* Petals ciliate along margins only……G. macbridei
26. Petals linear–ligulate…………………………...…27
26* Petals oblong–elliptic………………………….…28
27. Inflorescence laxly several-flowered, petals ciliolate
along margins…………………………..…G. bogotensis
27* Inflorescence densely many-flowered, petals ciliate
along margins…………………………….…G. caucana
28. Leaves arranged along stem, decreasing in size
upwards…………………………………….…G. foliosa
28* Leaves gathered in the basal rosette………….…29
29. Petals 3-nerved, lip apical part papillate along
margins, gynostemium pubescent just below stigma
towards the base…………………..…G. cundinamarcae
29* Petals 1-nerved, lip apical part densely ciliate along
margins, gynostemium ciliate in the lower half
only……………………………………………G. adnata
Taxonomic treatment
Gomphichis pseudogoodyeroides S. Nowak
and Szlach., sp. nov. (Fig. 1)
Gomphichis pseudogoodyeroides is related to G. goodyeroides Lindl., but the former have elliptic petals wider than
dorsal sepal and lip with prominent furrow along midnerve,
with two fleshy and thick keels along its margins.
Type: H. Garcia Barriga 17592—Colombia, Cundinamarca, Sasaima, San Bernardo, Quebrada La Maria y Rio
Dulce, elev. 1,750–1,950 m (20–30 Nov 1962), (COL!
holotype).
Plants up to 100 cm tall, erect, rather stout. Leaves
basal, petiolate; petiole to 8 cm long, rather wide; blade up
to 23 cm long and up to 3.5 cm wide; oblong–lanceolate or
linear–lanceolate, acute to subacute. Scape erect, several
sheathed, glabrous below, glandular above. Spike up to
15 cm long, cylindrical, densely many flowered. Flowers
pubescent externally especially towards the base. Floral
bracts up to 9 mm long, ovate–lanceolate, acute, pubescent. Ovary 5 mm long, sessile, ovoid, densely tomentose.
Dorsal sepal up to 6 mm long, 2.5 mm wide, concave,
oblong–elliptic, acuminate to shortly apiculate. Petals up to
5 mm long and 2.7 mm wide, elliptic with broad base and
acute apex, sessile, margins ciliolate except base and apex,
3-nerved. Lateral sepals up to 6 mm long and 3 mm wide,
obliquely ovate–elliptic, acute to acuminate, concave. Lip
up to 6 mm long in total, divided into hypochile and epichile; hypochile 4 mm long, 5 mm wide, trapezoid in
outline, thick and papillate in the centre, with two small
thickenings at the base; epichile 2 mm long, 1 mm wide,
oblong–elliptic, acute, with prominent furrow along midnerve, with two fleshy and thick keels along its margins.
Gynostemium 5 mm long, pubescent dorsally, ciliate
ventrally.
123
S. Nowak et al.
Fig. 2 Distribution map of Gomphichis pseudogoodyeroides S.
Nowak and Szlach. in Andean Colombia
Fig. 1 Gomphichis pseudogoodyeroides S. Nowak and Szlach.:
a dorsal sepal, b petal, c lateral sepal, d lip-side view, e lip,
f gynostemium [H. Garcia Barriga 17592 (COL); drawn by S. Nowak]
Ecology: Terrestrial in paramo, subparamo with Weinmannia tomentosa. Flowering time: January, March, June–
August, October–November.
Distribution: Colombia (Boyacá, Cesar, Cundinamarca,
Magdalena, Nariño, Norte de Santander, Quindio). Elev.
1,050–3,410 m (Fig. 2).
Etymology: An allusion to the similarity of Gomphichis
goodyeroides.
Conservation status: According to the IUCN Red List
(IUCN 2011), the species can be assigned as near threatened (NT). The species is known from several localities;
however all of them are limited to high Andean páramo,
which is heavily impacted (Davis et al. 1997), so there is
not much potential habitat. Therefore, in our opinion,
continuing decline in extent and area of occurrence is
expected.
Representative specimens: G. Huertes G. and L.
A. Camargo G. 6832—Colombia, Boyaca, Mpio. de Arcabuco, páramo, elev. 3,300 m (3 Jan 1970), (COL!); H.
Schmidt-Mumm 77—Colombia, Boyaca, Mpio. de
123
Arcabuca, entre Tunja y Arcabuco, elev. 2,900 m (6 Jun
1961), (COL!); D. Stancik, H. Mendoza and S. Medina
1202—Colombia, Boyaca, Mpio. Villa de Leyva, Vereda
La Capilla, Finca Manantiales de Iguaque, 58410 N,
738290 W, elev. 2,600 m (5 Nov 1998), (COL!); A. Etter
and L. G. Baptiste 907—Colombia, Boyaca, Mpio. de
Boavita, Cañon del Chicamocha, Bosque Chulavita, elev.
3,000 m (2 Oct 1991), (COL!); L. A. Camargo G. and G.
Huertas G. 7793—Colombia, Boyaca, Mpio. de Santa
Rosa de Viterbo, Cordillera Oriental, cerros orientales
vecinos a la poblacion, elev. 2,780 m (13 Jan 1981),
(COL!); A. M. Cleef 2240 —Colombia, Boyaca, páramos al
NW de Belen, subida al Alto de las Cruces, hacia San José
de la Montana, subpáramo seco 5 km NW de Belen, elev.
3,410 m (5 Mar 1972), (COL!); H. Garcia Barriga and R.
Jaramillo M. 20686—Colombia, Cesar/Norte de Santander, Cordillera Oriental, Jurisdicciones Cerro de Oroque,
elev. 3,000–3,900 m (22–27 Jul 1974), (COL!); D.R.
Jimenez and S. Cortés 130—Colombia, Cundinamarca,
Mpio. de Tausa, Vereda Lagunitas, Cerro San Isidro, elev.
3,140 m (8 Jul 1998), (COL!); S. Hernandez M. 366 —
Colombia, Cundinamarca, Mpio. de Subachoque, Finca El
Cerro (Vereda El Tobal), en un bosque predominantemente
de encenillos (Weinmannia tomentosa), sobre el suelo, en
un colchón de musgo, elev. 2,960 m (3 Nov 1998), (COL!);
L. Uribe U. 330—Colombia, Cundinamarca, Cerca a
Eight new species of Gomphichis
Zipaquirá, elev. 2,700 m (1939), (COL!); J.L. Fernandez
Alonso, H. Cardozo and M. Arevalo 11594—Colombia,
Cundinamarca, Bogotá-Chocontá, Represa del Sisga, Pasando el Puente Metálico, elev. 2,600 m (20 Aug 1994),
(COL!); M. Schmidt 128—Colombia, Cundinamarca, Zipacón, elev. 2,700 m, (COL!); L. Uribe U. 6001—Colombia, Cundinamarca, Suba, Colinas al oriente, vertiente
hacia la sabana de Bogotá, elev. 2,700 m (12 Nov 1967),
(COL!); H. Garcia Barriga 17592—Sabana de Bogota,
Chia, La Capilla, elev. 2,700 m, (COL!); R. Castañeda
7549—Colombia, Magdalena, Cordillera Oriental, Camino
de Codazzi a Machiques, antes de la frontera, elev.
2,400 m (11 Mar 1959), (COL!); L.E. Mora 2301—
Colombia, Narino, en la carretera entre Barbacoas y Junı́n,
elev. 1,050 m (7 Aug 1962), (COL!); Kapuler and Hascall
240—Colombia, Quindio, Sine loc., common species not
collected, i.e. those on mud wall, rather those from trees
included, (Aug 1964), (COL!).
Notes: One of the most common Gomphichis species in
Colombia. It is easily separable from its closest relative G.
goodyeroides by the lip and petals form. The petals of G.
pseudogoodyeroides are elliptic and wider (up to 2.7 mm)
than dorsal sepal (up to 2.5 mm), in contrast to G.
goodyeroides where petals are obovate and narrower (up to
2.2 mm) than dorsal sepal (up to 3 mm). Lip of the new
species is explicitly divided into hypochile and epichile
(vs. more or less constricted at the apex in G. goodyeroides), in the centre of basal half thick and papillate (vs. pair
of tomentose, cushion-like thickenings in the apical half).
Additionally, epichile part of G. pseudogoodyeroides lip
with prominent furrow along midnerve and two fleshy,
thick keels along its margins, which are absent in G.
goodyeroides.
Gomphichis salamancae S. Nowak and Szlach., sp.
nov. (Fig. 3)
Species related to Gomphichis goodyeroides, from which it
differs by sessile, elliptic or oblong–elliptic petals with
broad base, and lip with massive cushion-like thickening in
the apical half, which is sulcate in front, gynostemium
ciliate on both surfaces.
Type: H. Garcia Barriga 20219—Colombia, Cundinamarca, Guasca, Paramo de Guasca, elev. 2,800 m (19 Nov
1971), (COL! holotype).
Plants up to 70 cm tall, erect, slender. Leaves three basal
and one cauline, petiolate; petiole up to 12 cm long, rather
wide; blade up to 16 cm long and up to 2.5 cm wide,
oblong–lanceolate or linear–lanceolate, acute. Scape erect,
several sheathed, glabrous below, glandular above. Spike
up to 6 cm long, cylindrical, densely many flowered.
Flowers pubescent externally especially towards the base.
Floral bracts up to 12 mm long, ovate–lanceolate, acute,
Fig. 3 Gomphichis salamancae S. Nowak and Szlach.: a dorsal
sepal, b petal, c lateral sepal, d lip-side view, e lip, f gynostemium [H.
Garcia Barriga 20219 (COL); drawn by S. Nowak]
pubescent. Ovary 5 mm long, sessile, ovoid, densely
pubescent. Dorsal sepal up to 6 mm long, 2.7 mm wide,
concave, oblong–elliptic, subacute. Petals up to 5 mm long
and 3 mm wide, elliptic (or oblong–elliptic) with broad
base and subacute apex, sessile, somewhat oblique, margins ciliolate except base and apex, 3-nerved. Lateral
sepals up to 6.5 mm long and 3 mm wide, obliquely ovate–
elliptic, subobtuse, concave. Lip up to 6.5 mm long in
total, divided into hypochile and epichile; hypochile 4 mm
long, 7 mm wide, transversely elliptic in outline, lateral
lobes truncate, with two small thickenings at the base;
epichile 2.5 mm long, 2 mm wide, elliptic, acute, with
massive cushion-like thickening, which is sulcate in front.
Gynostemium 5 mm long, ciliate on both surfaces.
Ecology: Terrestrial in páramo, subpáramo with
Weinmmania tomentosa, Diplostephium, Myrsine and
Cavendishia. Flowering time: February, September,
November.
Distribution: Colombia (Cundinamarca, Norte de Santander, Risaralda). Elev. 400–3,000 m (Fig. 4).
Etymology: Dedicated to Sonia Salamanca Villegas, in
the past a very active collector of Colombian plants and
Andean vegetation plants.
Conservation status: According to the IUCN Red List
(IUCN 2011), Gomphichis salamancae can be assigned as
123
S. Nowak et al.
as G. pseudogoodyeroides, but in contrast to G. goodyeroides where petals are obovate and narrower than dorsal
sepal. Epichile of the lip of the new species with massive
cushion-like thickening, which is sulcate in front (vs.
prominent furrow along midnerve and two fleshy, thick
keels along its margins in G. pseudogoodyeroides and
absence of such structures in G. goodyeroides). Additionally, gynostemium of G. salamancae is ciliate on both
surfaces (vs. pubescent dorsally, ciliate ventrally in G.
pseudogoodyeroides and G. goodyeroides).
Gomphichis carlos-parrae S. Nowak and Szlach., sp.
nov. (Fig. 5)
Fig. 4 Distribution map of Gomphichis salamancae S. Nowak and
Szlach. (circle) and G. carlos-parrae S. Nowak and Szlach. (triangle)
in Andean Colombia
vulnerable (VU B1ab(i,iii)). The species is known only
from five localities. An extent of its occurrence is about
39,000 km2, but all of locations are restricted to high
Andean páramo, which is heavily impacted (Davis et al.
1997) and it may be a reason that real EOO (suitable
habitat for species) is much narrower.
Representative specimens: M. Hernandez Schmidt
1034—Colombia, Cundinamarca, Mpio. de Subachoque,
Vereda Tobál, finca El Cerro, en un bosque de porte bajo
dominado por Weinmannia tomentosa, y con presencia de
Diplostephium, Myrsine y Cavendishia, (11 Nov 2002),
(COL!); M. Schneider 692—Colombia, Cundinamarca, El
Retiro, elev. 2,950 m (27 Sep 1953), (COL!); J. L. Fernández Alonso, C. J. Orozco and P. Galvis 11845—
Colombia, Norte de Santander, Via Toledo hacia vereda
Sta Isabel, subida de la Finca Palo Colorado (hacia el
subpáramo) al Páramo de Santa Isabel, elev.
3,000–3,800 m (3 Nov 1994), (COL!); J. H. Torres, O.
Rangel, P. Franco, A. M. Cleef and S. Salamanca 217—
Colombia, Risaralda, Mpio. de Santuario, Vereda Las
Colonias, 400 m arriba del campamento, elev. 2,910 m (2
Feb 1983), (COL!).
Notes: It is related to Gomphichis goodyeroides and G.
pseudogoodyeroides from which it differs in the lip and
petals. The petals of G. salamancae are elliptic and wider
(up to 3 mm) than dorsal sepal (up to 2.7 mm) as the same
123
Gomphichis carlos-parrae has lip very similar to G. valida
Rchb.f., but differs from the latter in having very long
leaves, petals as long as dorsal sepal and gynostemium
strongly swollen above narrow base and densely ciliate
below stigma.
Type: Langenheim 3475—Colombia, Boyaca, near
Buenos Aires (Station 95) in Páramo de La Rusia on road
between Duitama and Charalá, ecotone between cloud
forest and paramo, elev. 3,650 m (17 Aug 1953), (COL!
holotype).
Fig. 5 Gomphichis carlos-parrae S. Nowak and Szlach.: a dorsal
sepal, b petal, c lateral sepal, d lip-side view, e lip, f gynostemium [J.
Langenheim 3475 (COL); drawn by S. Nowak]
Eight new species of Gomphichis
Plants up to 110 cm tall, robust. Leaves numerous, basal
and few cauline, without prominent petiole, up to 40 cm
long and 1.5 cm wide, linear–lanceolate, acute, erect,
tapering towards vaginate base. Scape erect, several
sheathed, terminated densely many-flowered cylindrical
spike. Flowers rather large, densely hirsute externally.
Floral bracts up to 15 mm long, oblong–elliptic, acute,
sparsely glandular. Ovary up to 10 mm long, cylindrical to
fusiform, pubescent. Dorsal sepal up to 8.5 mm long and
4.6 mm wide, ovate–oblong, obtuse, 3-nerved. Petals up to
8.5 mm long and 4.1 mm wide, from a cuneate base obliquely elliptic–obovate or obovate, attenuate towards base,
shortly petiolate, externally as well as along the margins
subdensely pubescent along the expanded portion, otherwise glabrous or sparsely pubescent on the outer surface,
l-nerved. Lateral sepals up to 9 mm long and 5 mm wide,
obliquely elliptic–oblong, obtuse, 3-nerved. Lip up to
10 mm long and 8 mm wide, from a cuneate base broadly
obovate in outline, sessile, widest at base, apical half with
two cushion-like thickening separated by furrow, apex
foveolate, base with two, thickenings. Gynostemium
3.5 mm long, much swollen above narrow base, ciliate
below stigma.
Ecology: Terrestrial in páramo, grassy páramo with
Calamagrostis effusa, woodland with Weinmannia microphylla and Clethra bicolor. Flowering time: January–
December.
Distribution: Colombia (Boyacá, Caldas, Cauca, Chocó,
Cundinamarca, Huila, Nariño, Putumayo, Risaralda). Elev.
2,850–3,800 m (Fig. 4).
Etymology: Dedicated to Dr. Carlos Parra, the Curator
of the National Herbarium of Colombia, Bogota.
Conservation status: According to the IUCN Red List
(IUCN 2011), the species can be assigned as near threatened (NT). The species is known from several localities,
however all of them are limited to high Andean páramo
similarly to Gomphichis pseudogoodyeroides, but G. carlos-parrae is characterized by much more restricted elevation requirements.
Representative specimens: J. Farfan s.n.—Colombia,
Boyaca, Mpio. de Chinavita, Cerro Mamapacha, Vereda
Mundo Nuevo, elev. 3,000 m (27 Jul 2001), (COL!); O.
Rangel 11660 et al.—Colombia, Boyaca, Mpio. Duitama.
Vereda El Carmen, El Rosal. Cerca a la Recebra, bosque de
Weinmannia microphylla y Clethra bicolor, elev. 3,390 m
(17 Mar 1994), (COL!); Stancik 1670—Colombia, Boyaca,
Mpio. de Sotaquirá, Páramo Chontales, shrubby–grassy
páramo on the summit (S slopes) of the ridge, elev.
3,100 m (15 Dec 1998), (COL!); A. M. Cleef 9423—
Colombia, Boyaca, Peña de Arnical, N de Vado Hondo,
lajas de arenisca del lado SE, elev. 3,470 m (6 Apr 1973),
(COL!); R. Castañeda 3025—Colombia, Caldas, Navarco,
elev. 3,000 m (12 Sep 1951), (COL!); G. Reina, L. Pito, S.
Tombe, R. Hurtado, A. Chilo and Chilo 447 and 447a —
Colombia, Cauca, Mpio. Silvia, Pitayo, sobre la via que del
crucero de Asnenga conduce Mosoco, area Especial de
Manejo Páramo de Moras, Vegetación subpáramo y páramo, elev. 3,000–3,600 m (4 Mar 2000), (COL!); M.P.
Córdoba and L. Bernal 2432—Colombia, Cauca, Mpio. de
Toribio, Corregimiento de Tacueyo, Páramo de Santo
Domingo,
Cordillera
los
Alpes,
28570 47.8200 N,
0
00
76807 57.22 W, elev. 3,495 m (16 Dec 2002), (COL!);
Downer 104—Colombia, Cauca, Macizo Central Colombiano, Alto del Buey, Farallones, elev. 3,600–3,800 m (22
Feb 1979), (COL!); J. Cuatrecasas 18926—Colombia,
Cauca, Cordillera Central, vertiente occidental, Cabeceras
del Rio Palo, quebrada del Rio Lopez, Quebrada del
Duende, elev. 3,400–3,450 m (2 Dec 1944), (COL!); C.
Cristancho 956—Colombia, Cauca, transecto cabaña INDERENA-Páramo de Letreros-Laguna de Santiago (Mirador), elev. 3,200–3,640 m (15 Feb 1987), (COL!); J.
Idrobo, P. Pinto and H. Bischler 3105—Colombia, Cauca/
Huila, Macizo Colombiano, Páramo de Las Papas, Colinas
al SE de la Laguna La Magdalena sobre el cerro La Corona
y El Boquerón, vertiente Magdalena, elev. 3,330–3,520 m
(8 Sep 1958), (COL!); J. H. Torres, O. Rangel, P. Franco,
A. M. Cleef and S. Salamanca 1824—Colombia, Choco,
Macizo del Tamaná, bajando del paso al valle de las
Mirlas, elev. 3,700 m (10 Feb 1983), (COL!); J. L. Fernandez Alonso, J. Groenendijk, D. Cortés and G. Peñaloza
19054—Colombia, Cundinamarca, Mpio. de SuescaNemocón, Hacienda Susatá, zona seca con restos de bosque
andino y subpáramo, elev. 2,850–2,950 m (23 Aug 2000),
(COL!); Y. Figueroa C. and N. Garcia 582—Colombia,
Cundinamarca, Bogotá, Localidad 5, Usme, sector del
Embalse de Chisacá, alrededores del rio Tunjuelo, Vereda
las Margaritas, 48200 N, 748150 W, elev. 3,000–3,200 m (19
Jun 2005), (COL!); Y. Figueroa C., J. Mora, T. Vivas and
O. Vargas 804—Colombia, Cundinamarca, Bogotá, Localidad 5, Usme, sector del Embalse de Chisacá, Vereda
Las Margaritas, 48200 N, 748150 W, elev. 3,000–3,200 m (6
Oct 2005), (COL!); P. Franco, J. Betancur and A. Neira
6146—Colombia, Cundinamarca, Santafé de Bogotá DC,
Sumapaz, corregimiento San Juan, vereda Capitolio, finca
La Pradera, 48170 N, 748120 W, elev. 3,150–3,200 m (27 Feb
1999), (COL!); O. Haught 5758—Colombia, Cundinamarca, road to E of Guasca, elev. 3,100 m (23 May 1947),
(COL!); Da Ros s.n.—Colombia, Cundinamarca, Guavio,
Guasca, Páramo de Guasca, Carretera de Bogotá, elev.
3,100 m (3 Jan 2001), (COL!); M.P. Córdoba, C. Lopez
and L. Bernal 2973—Colombia, Cundinamarca, Mpio. de
Chocontá, 5840 58.700 N, 738440 13.700 W, elev. 2,850 m (29
Nov 2003), (COL!); B. Cesar 9431—Colombia, Huila, Al
Alto de la Linea, 28470 0700 N, 768010 2600 W (4 Dec 1993),
(COL!); B. Cesar 9449—Colombia, Huila, Al Alto de la
Linea, 28470 0700 N, 768010 2600 W (6 Dec 1993), (COL!); A.
123
S. Nowak et al.
Fernandez and K. E. Knoth 1025—Colombia, Nariño, a lo
largo de la carretera de Pasto a Sibundoy, elev.
2,600–2,800 m (5 Jan 1952), (COL!); D. Stancik 2795—
Colombia, Nariño, Mpio. Túquerres, Volcán Azufral, road
from the vereda San Roque Alto to Laguna Verde, km 6,
grassy páramo (Calamagrostis effusa, Cortaderia, Diplostephium), elev. 3,800 m (5–9 Mar 1999), (COL!); A.
Muñoz and B. Ramirez 312—Colombia, Nariño/Putumayo,
Mpio. de Santiago, Vereda San Antonio de Bellavista,
Páramo del Bordoncillo, 18110 N, 778060 W, elev.
3,200–3,400 m (17 Apr 1993), (COL!); J. Cuatrecasas
11729—Colombia, Putumayo, Alta Cuenca del Rio Putumayo, filo de la Cordillera entre El Encano y Sibundoy,
Páramo de San Antonio del Bordoncillo, 18110 N, 778060 W,
elev. 3,250 m (4 Jan 1941), (COL!); D. Stancik 3427—
Colombia, Risaralda, Mpio. de Santuario, NP Tatamá,
patches of the grassy paramo (Calamagrostis effusa)
between of the matorral, elev. 3,750 m (23 Sep 1999),
(COL!); W.G. Vargas 8676—Colombia, Tolima, Mpio. de
Cajamarca, Corregimiento de Anaime, Páramo de Valles,
elev. 3,500 m (Dec 2000), (COL!); J. Cuatrecasas and
Echeverry 27657—Colombia, Tolima, Cordillera Central,
La Linea, cerro El Campanario, subpáramo y páramo, elev.
3,600–3,700 m (4 Mar 1969), (COL!).
Notes: Gomphichis carlos-parrae is related to G. valida,
lip of both species is very similar, but the former species
possesses much longer leaves up to 40 cm (vs. 9 cm in G.
valida). The petals of G. carlos-parrae are as long as dorsal
sepal (up to 8.5 mm), in contrast to G. valida where petals
(up to 6.7 mm) are shorter than dorsal sepal (up to 7 mm).
In the new species gynostemium is shorter and massive,
also densely ciliate below stigma (vs. pubescent below
stigma in G. valida).
Gomphichis magnicallosa S. Nowak and Szlach., sp.
nov. (Fig. 6)
Plants relatively small, delicate, somewhat similar to
Gomphichis traceyae Rolfe, but with petals pubescent
along margins and very large, prominent oblong basal
thickenings being 1/3 of the whole lip length.
Type: J. L. Fernandez Alonso, M. R. Garzón et al.
7745—Colombia, Cundinamarca, DC de Bogota, Páramo
Sumapaz, después de la laguna de Chisacá, bosque andino
y pajonales cerca del rı́o Taquecito, elev. 3,500–3,700 m
(9–13 Nov 1987), (COL! holotype).
Plants 35–40 cm tall, delicate, erect. Leaves 5–6, basal,
shortly petiolate; petiole 1–2 cm long; blade to 5 cm long,
1.7 cm wide, oblong–lanceolate to oblong–elliptic, acute.
Scape several sheathed, glabrous in the lower part, glandular above. Flowers small, densely pubescent externally.
Floral bracts up to 8 mm long, lanceolate, acute, sparsely
glandular. Ovary 4–5 mm long, cylindrical, pubescent.
123
Fig. 6 Gomphichis magnicallosa S. Nowak and Szlach.: a dorsal
sepal, b petal, c lateral sepal, d lip-side view, e lip, f gynostemium [J.
L. Fernandez Alonso, M. R. Garzón et al. 7745 (COL); drawn by S.
Nowak]
Dorsal sepal up to 4 mm long and 1.8 mm wide, obovate–
oblong, obtuse, 1-nerved. Petals up to 4 mm long and
1.5 mm wide, from a cuneate base obliquely oblong–
obovate or obovate, attenuate towards base, elongate
towards truncate apex, pubescent along margins, except
apex, obscurely 3-nerved. Lateral sepals up to 4 mm long
and 1.8 mm wide, obliquely elliptic–oblong, obtuse,
1-nerved. Lip up to 4.2 mm long and 3 mm wide, with
very large, massive, prominent oblong basal thickenings
being 1/3 of the whole lip length, hypochile 3 mm long,
trapezoid in outline, pubescent in the centre; epichile
1 mm long and wide, ligulate–ovate, subobtuse, sulcate.
Gynostemium 3.8 mm long, minutely ciliate all over
below stigma.
Ecology: Terrestrial mainly in grassy páramo dominated
by Espeletia, Calamagrostis, Hypericum, Lupinus, Monnina, Cavendishia, and Castilleja, also shrubby páramo,
mountain forest with Prumnopitys and Podocarpus.
Flowering time: April–May, July–December.
Distribution: Colombia (Boyacá, Cauca, Cesar, Cundinamarca, Magdalena). Elev. 2,600–3,800 m (Fig. 7).
Eight new species of Gomphichis
Etymology: In reference to the presence of large basal
lip calli.
Conservation status: According to the IUCN Red List
(IUCN 2011), the species can be assigned as NT. The
species is known from several locations and occurs at the
highest elevations at which the representatives of Gomphichis have been reported ever.
Representative specimens: N. Diaz 32—Colombia,
Boyaca, Mpio. de Villa de Leyva, Santuario de Flora y
Fauna de Iguaque, camino a la laguna, elev. 3,800 m (14
Nov 1988), (COL!); H. Dueñas, F. Cortés and N.
Aranguren 3129—Colombia, Boyaca, Mpio. de Arcabuco,
Santuario de Flora y Fauna de Iguaque, camino a la
laguna, paramo dominado por Espeletia, Calamagrostis,
Hypericum, Lupinus, Monnina, Cavendishia, Castilleja,
elev. 3,600 m (24 Sep 2002), (COL!); M.P. Córdoba, C.
Lopez and L. Bernal 3038—Colombia, Boyaca, Mpio. de
Tibasosa, Cerro Guaquita, 58420 33.200 N, 738020 16.300 W,
elev. 3,120 m (11 Dec 2003), (COL!); J. Farfan s.n.—
Colombia, Boyaca, Mpio. de Tunja, Rio Teatinos, Embalse de Teatinos, bosque, elev. 2,800 m (10 Oct 2001),
(COL!); L.A. Camargo G. 8048—Colombia, Boyaca,
Mpio. de Socotá, Cordillera Oriental, en matorrales, a la
orilla de la carretera que conduce al Páramo de Pisba,
elev. 2,950 m (30 Sep 1981), (COL!); D. Stancik 1304—
Colombia, Boyaca, Mpio. de Samacá, Vereda Ruch cal––
Fig. 7 Distribution map of Gomphichis magnicallosa S. Nowak and
Szlach. in Andean Colombia
parte alta, shrubby matorral on the margin of road from
Vereda to Desaquadero, elev. 2,900 m (1 Nov 1998),
(COL!); P. J. Grubb, B. A. B. Curry and A. Fernández
Pérez 609—Colombia, Boyaca, Sierra Nevada del Cocuy,
on ground in hill sabana (?forest cleared) between Laguna
Seca and Bachira by path, elev. 2,750 m (19 Aug 1957),
(COL!); P.J. Grubb, B.A.B. Curry and A. Fernández
Pérez 773—Colombia, Boyaca, Sierra Nevada del Cocuy,
on ground near river in shrub paramo zone near El
Playón, elev. 3,800 m (10 Sep 1957), (COL!); F.A.Barclay and R.E. Schultes 144—Colombia, Cauca, Mpio. de
Puracé, N slope of the Volcán de Puracé, elev.
2,700–2,800 m (23 Jul 1956), (COL!); H. Barclay and P.
Juajibioy 5910—Colombia, Cauca, Macizo Colombiano,
Valle de Las Papas, ver wet portion of extensive Espeletia-grass cienaga near Hacienda Los Andes, Sta
4.2–2.5 km from casa Los Andes, elev. 3,150 m (4 Oct
1958), (COL!); E.P. Arbelaez and J. Cuatrecasas 5941—
Colombia, Cauca, Del Páramo a Puracé, Matorrales en
Chiquin, elev. 2,700–3,100 m (11 Jul 1939), (COL!); G.
Lozano C. 3509—Colombia, Cauca, Mpio. de Coconuco,
páramo del valle de Paletara, elev. 2,950 m (30 Jul 1980),
(COL!); O. Rangel, P. Franco, A. Rudas, J.R. Olmos, M.
Pardo and M. Clavijo 11047b—Colombia, Cesar, Mpio
de Manaure, Serrania del Perijá, Casa de Vidrio, 32 km
SE de Manaure, Arriba de la Laguna, en bosque de
Prumnopytis y Podocarpus, 728530 W, 108250 N, elev.
2,950 m (5 Nov 1993), (COL!); C. Hahn and K. E. Koch
s.n.—Colombia, Cundinamarca, Mpio. de Bogotá, entre
Bogotá y Choachı́, Páramo de Cruz Verde, cerca del camino al cerro de antenna de television, cerca al barranca
del camino (24 Apr. 1996), (COL!); J.L. Fernández
Alonso, M.R. Garzón et al. 7745—Colombia, Cundinamarca, DC de Bogotá, Páramo Sumapaz, después de la
laguna de Chisacá, bosque andino y pajonales cerca del
rio Taquecito, elev. 3,500–3,700 m (9–13 Nov 1987),
(COL!); P. Pedro, P. Franco, D. Stancik and A. Neira
216—Colombia, Cundinamarca, DC, localidad 20, Parque
Nacional Natural Sumapaz. Vereda Santa Rosa, Ladera de
la Quebrada Bijuacales (7 Aug 1998), (COL!); M.R.
Garzón 178—Colombia, Cundinamarca, Bogota, Páramo
de Monserrate, Vereda El Verjón, Hacienda Santa Barbara, El Granizo, elev. 3,000–3,200 m (15–23 Nov 1986),
(COL!); M.R. Garzón 337—Colombia, Cundinamarca,
Bogota, Páramo de Monserrate, Vereda El Verjón, Hacienda Santa Barbara, El Granizo, elev. 3,000–3,200 m (3
Oct 1987), (COL!); M. Schneider 245/1—Colombia,
Cundinamarca, Bogota, Cerro de Monserrate, elev.
3,100 m (17 Sep 1944), (COL!); M. Schneider 245/2—
Colombia, Cundinamarca, Bogota, Cerro de Monserrate,
elev. 3,100 m (17 Sep 1944), (COL!); M. Schneider
288/1—Colombia, Cundinamarca, Bogota, Cerro de
Monserrate, elev. 3,100 m (17 Sep 1944), (COL!); M.
123
S. Nowak et al.
Schneider 288/3—Colombia, Cundinamarca, Bogota,
Cerro de Monserrate, elev. 3,250 m (2 Jul 1950), (COL!);
J. Cuatrecasas 20—Colombia, Cundinamarca, Bogota,
Cerro de Monserrate, Cumbre, matorroles, elev. 3,210 m
(20 Aug 1938), (COL!); M.A. Bello 86—Colombia,
Cundinamarca, Mpio. La Calera, Páramo Chingaza,
Monteredondo, camino a La Quebrada Babilonia,
48380 200 N, 738430 800 W, elev. 3,000 m (27 Dec 1999),
(COL!); S. P. Cortés Sánchez 555—Colombia, Cundinamarca, Mpio. de Chia, Cerca de la casa del cabildo indijena, elev. 2,690 m (17 May 1996), (COL!); M.
Schneider 245/3—Colombia, Cundinamarca, Usme, elev.
2,600–2,700 m (26 Sep 1951), (COL!); G. Morales L.
37—Colombia, Cundinamarca, Usme, embalse de Chisacá, elev. 3,100 m (27 Oct 1977), (COL!); M. Schneider
288/4—Colombia, Cundinamarca, Usaquén, elev. 3,000 m
(12 Oct 1951), (COL!); L. Uribe U. 5998—Colombia,
Cundinamarca, Tocancipá, montes altos al oriente, elev.
2,800 m (5 Nov 1967), (COL!); Haught 6492—Colombia,
Cundinamarca, Guadelupe near Bogotá, Open páramo,
elev. 3,300 m (17 Jul 1949), (COL!); G. Huertas and L.
A. Camargo G. 6573—Colombia, Cundinamarca, Mpio.
de Fómeque, Páramo de Chingaza, Cerros vecinos a La
Laguna, elev. 3,400 m (3 Nov 1966), (COL!); G. Morales
L. and Flenley 31—Colombia, Cundinamarca, Páramo
Cruz Verde, elev. 3,200 m (6 Sep 1977), (COL!); M.
T. Murillo 400—Colombia, Cundinamarca, Zipaquirá, El
Carrizal, elev. 3,200–3,400 m (14 Oct 1961), (COL!);
Barclay 5343—Colombia, Cundinamarca, Páramo de
Guerrero, between Zipaquirá and Pacho, elev. 3,200 m (9
Jul 1957), (COL!); B. van Wesenbeeck and J. van Mourik
6—Colombia, Cundinamarca, Tominé de Indios, bosque
altoandino y subparamo, elev. 3,000–3,100 m (3 Aug
1999), (COL!); J.L. Fernandez Alonso, O. Rangel, S.
Cortés, A. Garzón and M. Hernandez 15556—Colombia,
Cundinamarca, Villapinzón, de Villapinzón a Umbita, limite municipal, elev. 3,200 m (21 May 1998), (COL!); J.
Cuatrecasas and R. Romero Castañeda 25060—Colombia, Magdalena, Sierra de Perijá, E of Manaure, Sabana
Rubia, páramo, elev. 3,000–3,100 m (6–8 Nov 1959),
(COL!).
Notes: Gomphichis magnicallosa is related to G.
traceyae, but its smaller plant. The new species is
34–40 cm tall (vs. up to 60 cm tall in G. traceyae), leaves
are petiolate, 6–7 cm long, 1.7 cm wide (vs. leaves sessile,
up to 10 cm long, 2.5 cm wide), lip up to 4.2 mm long and
3 mm wide (vs. up to 4.5 mm long and 5 mm wide).
Additionally lip of G. magnicallosa with very large, massive, prominent oblong basal thickenings being 1/3 of the
whole lip length, what is unusual in the genus. Gynostemium of G. magnicallosa is minutely ciliate all over below
stigma, in contrast to G. traceyae where is almost glabrous
or glabrous.
123
Gomphichis fernandezii S. Nowak and Szlach., sp.
nov. (Fig. 8)
The species allied to Gomphichis traceyae, but with
1-nerved petals and lip with furrow along the midnerve and
thin apical part of epichile.
Type: J. L. Fernandez Alonso et al. 11921—Colombia,
Boyaca, Mpio. de Duitama, Trayecto Vereda El Carmen
(3,100 m) a el Páramo de La Rusia (3,550 m), (19 Nov
1994), (COL! holotype).
Plants up to 45 cm tall. Leaves 2–3 basal and 1–2
cauline, petiolate; petiole up to 6 cm long; blade up to
5 cm long and 1.5 cm wide, oblanceolate to oblong–lanceolate, acute. Scape slender, erect, glabrous in the lower
part, glandular towards the apex, with seven cauline
sheaths. Flowers small. Floral bracts up to 9 mm long,
ovate–lanceolate, acute, sparsely pubescent. Ovary up to
7 mm long, sessile, ovoid, glandular. Dorsal sepal up to
4 mm long and 1.8 mm wide, oblong–obovate, deeply
concave, subacute to subobtuse, glandular pubescent outside. Petals up to 3.8 mm long and 1.2 mm wide, obliquely
ovate–oblong or ovate–elliptic, obtuse, long-pubescent
along inner margin and ciliate along outer margins,
1-nerved. Lateral sepals up to 4 mm long and 2 mm wide,
Fig. 8 Gomphichis fernandezii S. Nowak and Szlach.: a dorsal sepal,
b petal, c lateral sepal, d lip-side view, e lip, f gynostemium [J.
L. Fernandez Alonso et al. 11921 (COL); drawn by S. Nowak]
Eight new species of Gomphichis
Fig. 9 Distribution map of Gomphichis fernandezii S. Nowak and
Szlach. (circle), G. monstruosa S. Nowak and Szlach. (square) and G.
lozanoi S. Nowak and Szlach. (triangle) in Andean Colombia
Fig. 10 Gomphichis jaramilloi S. Nowak and Szlach.: a dorsal sepal,
b petal, c lateral sepal, d lip-side view, e lip, f gynostemium [J.
Cuatrecasas and R. Jaramillo 12035 (COL); drawn by S. Nowak]
obliquely oblong–ovate, concave, subobtuse, glandular
pubescent outside. Lip up to 3.6 mm long in total, sessile,
fleshy, concave, difficult to spread, when spread divided
into hypochile and epichile; hypochile 2 mm long, 2.5 mm
wide, trapezoid–elliptic in outline; epichile 1.2 mm long,
0.8 mm wide, ligulate, with a pair of cushion-like calli,
separated by a furrow, apex of epichile thin. Gynostemium
2.5 mm long, minutely ciliate below stigma.
Ecology: Terrestrial in páramo with Polylepis, Espeletia, Melastomataceae, Ericaceae and Asteraceae. Flowering
time: February, November.
Distribution: Colombia (Boyacá, Cundinamarca). Elev.
3,400–3,600 m (Fig. 9).
Etymology: Dedicated to J. L. Fernandez Alonso, who
collected the specimens of this species.
Conservation status: According to the IUCN Red List
(IUCN 2011), the species can be assigned as endangered
(EN B1ab(i,iii)). The species is known only from three
localities. The known extent of occurrence of this species is
about 1000 km2 and the known localities are all restricted
to very narrow elevation range in high Andean paramo.
Representative specimens: J. Betancur et al. 5632—
Colombia, Boyaca, Mpio. de Duitama, Corregimiento El
Carmen, via a Virolin, Páramo de La Rusia, páramo muy
perturbado dominado por Polylepis, Espeletia, compuestas,
Melastomataceae y Ericaeae, elev. 3,400–3,500 m (19 Nov
1994), (COL!); M. Schneider 288/5—Colombia, Cundinamarca, Páramo de Siberia, elev. 3,600 m (24 Feb 1952),
(COL!).
Notes. The species appears to be related to Gomphichis
traceyae, from which it is easily separable by the petals and
lip form. The petals of G. fernandezii are long-pubescent
along inner margin and ciliate along outer margins, in
contrast to G. traceyae where petals are sparsely ciliolate
along margin in middle as well as on the back and
obscurely erose-denticulate at rounded apex. Additionally
lip of G. fernandezii with a pair of cushion-like calli,
separated by a furrow (vs. absent in G. traceyae) and thin
apex of epichile (vs. a pair of cushion-like, approximate
calli which are papillose). Gynostemium of G. fernandezii
is minutely ciliate below stigma, in contrast to G. traceyae
where is almost glabrous or glabrous.
Gomphichis jaramilloi S. Nowak and Szlach., sp. nov.
(Fig. 10)
This species is characteristic by its sessile rhombic lip, very
wide hypochile, subcordate epichile, ciliate petals and
123
S. Nowak et al.
Fig. 11 Distribution map of Gomphichis jaramilloi S. Nowak and
Szlach. in Andean Colombia
geniculate gynostemium, which is densely ciliate on the
dorsal surface and sparsely ciliate on the ventral one.
Type: J. Cuatrecasas and R. Jaramillo 12035—
Colombia, Cundinamarca, Cordillera Oriental, extremo SE
de la Sabana de Bogotá, en San Miguel, bosque, elev.
2,800–3,000 m (10 Sep 1941), (COL! holotype).
Plants ca. 75 cm tall. Leaves 1, basal, withering at
flowering. Scape with 7, glabrous sheaths, glabrous below,
glandular above. Spike 11.5 cm long, cylindrical, densely
many flowered. Flowers small. Floral bracts 7 mm, linear–
lanceolate, somewhat oblique. Ovary 10 mm long, densely
glandular. Dorsal sepal up to 4.5 mm long and 1.8 mm
wide, oblong–obovate, concave, subacute, densely pubescent outside, obscurely 1-nerved. Petals up to 4.5 mm long
and 1.7 mm wide, obliquely oblong–oblanceolate, attenuate towards subacute apex and towards the base, ciliate on
both margins near the middle, obscurely 3-nerved. Lateral
sepals up to 4.5 mm long and 2.5 mm wide, obliquely
oblong–elliptic, concave, subobtuse, densely pubescent
outside, 1-nerved. Lip up to 5 mm long in total and 5.5 mm
wide, sessile, with no basal thickenings; hypochile rhombic
when spread, very thick and papillate at the apex, sulcate;
epichile 1 mm long and wide, subcordate, subacute, thick
at the base. Gynostemium 4 mm long, geniculate above the
base, very ciliate on dorsal surface, sparsely ciliate on the
ventral one.
123
Ecology: Terrestrial in páramo, subpáramo, also Andean
forest with Weinmannia, Brunellia, Clusia and Miconia.
Flowering time: February–May, July–September, November.
Distribution: Colombia (Cundinamarca, Huila, Magdalena, Risaralda, Tolima), Venezuela (Mérida). Elev.
2,500–3,700 m (Fig. 11).
Etymology: Dedicated to Roberto Jaramillo Mejı́a, who
intensively collected and was an expert in Colombian
plants.
Conservation status: According to the IUCN Red List
(IUCN 2011), the species can be assigned as NT. The
species is known from several fragmented localities, most
of them are concentrated in Cundinamarca. All localities
are limited to heavily impacted high Andean páramo
(Davis et al. 1997), so there are not much potential habitat
and in our opinion continuing decline in extent and area of
occurrence, as well as a quality of habitat are expected.
Representative specimens: H. Dueñas and G. Peñalosa
3066—Colombia, Cundinamarca, Mpio. de Suesca-Nemocón, Vereda Susatá, Hacienda Susatá (4 May 2001),
(COL!); J.L. Fernandez Alonso, J. Groenendijk, D. Cortés
and G. Peñaloza 19054—Colombia, Cundinamarca, Mpio.
de Suesca-Nemocón, Hacienda Susatá, zona seca con restos de bosque andino y subpáramo, elev. 2,850–2,950 m
(23 Aug 2000), (COL!––sterile); H. Dueñas s.n.—Colombia, Cundinamarca, Mpio. de Bogotá, Jardı̂n Botánico
Bogota ‘‘José Celestino Mutis’’, elev. 2,650 m (20 Nov
2001), (COL!); M. Schneider 9/1—Colombia, Cundinamarca, alrededores de Bogotá, El Chicó, elev. 2,750 m (8
Apr 1944), (COL!); G. Lozano C., R. Castillo and E. Vega
728—Colombia, Cundinamarca, Carretera Bogotá-Choachı́, vertiente hacia Choachı́, adelante de Divorcium
Acuarum (10 Mar 1967), (COL!); S. P. Cortés Sánchez
816—Colombia, Cundinamarca, Mpio. de Chia, Cima del
cerro Manjuy, elev. 2,910 m (Aug 1996), (COL!––sterile);
M. Schneider 9/2—Colombia, Cundinamarca, Bojacá, elev.
2,800–2,900 m (28 Jun 1953), (COL!––sterile); G. Morales
L. 193—Colombia, Cundinamarca, Páramo de Guasca, km
48 via Sueva, elev. 3,100 m (25 Aug 1979), (COL!––in
bud); J.L. Fernandez Alonso, E. Linares, P. Balcazar, R.
Yasquez, J. Velez and G. Salazar 14884—Colombia, Huila,
Mpio. de Gigante, subida desde Vereda Ventanas al Páramo de Miraflores, bosque andino de Weinmannia, Brunellia, Clusia, Miconia, elev. 2,970 m (12–16 Aug 1997),
(COL!); R.Romero Castañeda 7751—Colombia, Magdalena, Mpio. de Santa Marta, entre Cerro Quemado y Cerro
San Lorenzo, elev. 2,600–2,800 m (16 Apr 1959), (COL!);
J.H. Torres, O. Rangel, P. Franco, A.M. Cleef and S.
Salamanca 2286—Colombia, Risaralda, Mpio. de Santuario, Vereda Las Colonias, margen derecha del rio San
Rafael, elev. 2,500 m (25 Feb 1983), (COL!); R. Echeverry
1990—Colombia, Tolima, camino de Nevada del Tolima,
Arriba del Rancho (Termales), elev. 3,700 m (17 Jul 1969),
Eight new species of Gomphichis
Fig. 12 Gomphichis monstruosa S. Nowak and Szlach.: a dorsal
sepal, b petal, c lateral sepal, d lip-side view, e lip, f gynostemium [G.
Lozano C. 3429 (COL); drawn by S. Nowak]
(COL!); H. Barclay and P. Juajibioy 9935—Venezuela,
Merida, alrededores de La Laguna Coromoto, Forestry
Station and Laguna Coromoto (2 Dec 1959), (COL!).
Notes: Gomphichis jaramilloi is easily separable from
its closest relative G. traceyae by the lip and gynostemium.
Lip of the new species as the same as G. traceyae is wider
than long, however differs with no basal thickenings (vs.
with small, oblong basal thickenings in G. traceyae).
Gynostemium of G. jaramilloi is very ciliate on dorsal
surface, sparsely ciliate on the ventral, in contrast to G.
traceyae where is almost glabrous or glabrous.
Gomphichis monstruosa S. Nowak and Szlach., sp.
nov. (Fig. 12)
Gomphichis monstruosa is related to G. longifolia (Rolfe)
Schltr. in having lip wider than long, but lip lateral lobes
are obliquely triangular, epichile with two thickenings near
the base and narrower, clawed petals.
Type: G. Lozano C. 3429—Colombia, Huila, Mpio. San
José de Isnos, Vereda El Marmol, Parque Nacional Naturel
de Puracé, elev. 2,800 m (24 Jul 1980), (COL! holotype).
Plants ca. 75 cm tall. Leaves 4, basal, sessile, to 16 cm
long and 1.7 cm wide, linear–oblanceolate, acuminate.
Scape with 5, sheaths, glabrous below, glandular above.
Spike 13 cm long, cylindrical, densely many flowered.
Flowers small. Floral bracts 12 mm long, linear–lanceolate,
somewhat oblique, pubescent. Ovary 7 mm long, densely
pubescent. Dorsal sepal up to 6 mm long and 3 mm wide,
oblong–obovate, concave, subobtuse, densely pubescent
outside, 1-nerved. Petals up to 6 mm long and 3 mm wide,
elliptic–ovate above cuneate base, attenuate towards acute
apex, somewhat oblique, ciliate on both margins near the
middle, otherwise glabrous, 1-nerved. Lateral sepals up to
5.7 mm long and 3.5 mm wide, obliquely oblong–elliptic,
concave, subobtuse, densely pubescent outside, 1-nerved.
Lip up to 6 mm long in total and 6.8 mm wide, clawed, with
two, basal thickenings; hypochile broadly rhombic when
spread, lateral lobes obliquely triangular, concave in the
lower half, thick and papillate at the apex, somewhat sulcate; epichile 1 mm long and wide, pentagonal, subacute,
with large cushion-like thickenings at the base distinctly
separated one from another. Gynostemium 4 mm long,
geniculate above the base, pubescent on dorsal surface,
ciliate on the ventral one.
Ecology: In forest. Flowering time: July, October.
Distribution: Colombia (Cundinamarca, Huila). Elev.
2,800 m (Fig. 9).
Etymology: An allusion to the lip which reminds, when
spread, a head of a monster.
Conservation status: According to the IUCN Red List
(IUCN 2011), the species can be assigned as vulnerable
(VU B1ab(iii)). The species is known only from two
fragmented localities and occurs at very high elevations, in
Andean forest, so this may be a reason for its rare occurrence. However, the distribution range of the species is
similar to the distribution range of Gomphichis lozanoi, so
in our opinion vulnerable category is appropriate.
Representative specimens: R. Echeverry 1990—
Colombia, Cundinamarca, bosque sobre Represa del Sisga,
tépalos verdosos, labello blanco amarillo (27 Oct 1984),
(COL!).
Notes: Gomphichis monstruosa is easily separable from
its closest relative G. longifolia by the lip and petals form.
The petals of G. monstruosa are elliptic–ovate above
cuneate base and longer (up to 6 mm) than lateral sepals
(up to 5.7 mm), in contrast to G. longifolia where petals are
obliquely obovate to broadly obovate and shorter (up to
5.5 mm) than lateral sepals (up to 6 mm). Additionally,
petals of G. monstruosa are ciliate on both margins near the
middle, otherwise glabrous (vs. ciliate all over margins in
G. longifolia). Lip of the new species as the same as G.
longifolia is wider than long, however epichile of G.
monstruosa with large cushion-like thickenings at the base
distinctly separated one from another differ from fleshy
callus in the central part of lip in G. longifolia.
Gomphichis lozanoi S. Nowak and Szlach., sp. nov.
(Fig. 13)
Species related to Gomphichis costaricensis (Schltr.)
Ames, F.T.Hubb. & C.Schweinf., but petals prominently
123
S. Nowak et al.
Fig. 13 Gomphichis lozanoi S. Nowak and Szlach.: a dorsal sepal,
b petal, c lateral sepal, d lip, e lip-side view, f gynostemium [G.
Lozano C. 3648 (COL); drawn by S. Nowak]
clawed, lip papillate in the centre and gynostemium bent at
right angel versus stigma. In G. costaricensis petals are
sessile, lip pubescent in the centre and gynostemium
geniculate above the base.
Type: G. Lozano C. 3648—Colombia, Cauca, Mpio. de
Tambo, La Romelia, Parque Nacional Natural Munchique,
elev. 2,600–2,800 m (5 Aug 1980), (COL! holotype).
Plants up to 120 cm tall. Leaves 2–5 basal and
occasionally 1 cauline, petiolate; petiole up to 10 cm
long, wide; blade to 22 cm long and 4 cm wide, linear–
oblanceolate to oblanceolate, acute. Scape with 4–7,
sheaths, slender, erect, delicate, glabrous below, glandular above. Spike 20–30 cm long, cylindrical, laxly
many flowered. Flowers realatively small. Floral bracts
8–10 mm long, linear–lanceolate, somewhat oblique,
pubescent on margins. Ovary 4–8 mm long, densely or
sparsely pubescent. Dorsal sepal up to 5.5 mm long and
2.8 mm wide, oblong–ovate, concave, subacute, pubescent outside, 1-nerved. Petals up to 5.2 mm long and
3.1 mm wide, elliptic–ovate to elliptic–suborbicular
above cuneate base, attenuate towards subacute apex,
oblique, ciliate on both margins near the middle,
otherwise glabrous, 3-nerved. Lateral sepals up to
5.5 mm long and 3 mm wide, obliquely oblong–elliptic,
123
concave, subobtuse, pubescent outside, 1-nerved. Lip up
to 5.7 mm long in total and 4.6 mm wide, clawed, with
two, basal thickenings; hypochile rhombic when spread,
lateral lobes obliquely triangular, concave in the lower
half, with two cushion-like, papillate thickenings, divided by prominent furrow; epichile 1 mm long and wide,
ligulate–ovate, subobtuse. Gynostemium up to 4.5 mm
long, bent at right angle above the base, ciliate below
stigma.
Ecology: Terrestrial in oak forest, also in páramo.
Flowering time: August, October–January.
Distribution: Colombia (Antioquia, Cauca, Chocó,
Cundinamarca, Norte de Santander). Elev. 2,300–2,900 m
(Fig. 9).
Etymology: Dedicated to G. Lozano C., who collected
type specimen of this species.
Conservation status: According to the IUCN Red List
(IUCN 2011), the species can be assigned as vulnerable
(VU B1ab(i,iii)). The species is known only from six
fragmented localities. An extent of occurrence is about
100.000 km2, but an occurrence of this species is
restricted mainly to oak forest, which is rare formation
in Andes, especially at high elevations, additionally the
oak forests are one of the less protected ecosystems
(Armenteras et al. 2003). All of these factors caused
that real EOO (suitable habitat for species) is probably
much narrower.
Representative specimens: G. Galeano et al. 304—
Colombia, Antioquia, Mpio. de Medellı́n, Vereda Santa
Elena, 12 km al E de Medellı́n, elev. 2,400 m (18 Dec
1980), (COL!); D. Stancik 1174—Colombia, Boyaca,
Mpio. de Gachantivá, Vereda Savedras de Roncancio,
Quercus-Clusia forest, terrestrial, 58430 3300 N, 738320 0100 W,
elev. 2,600 m (2 Nov 1998), (COL!); A. Gentry and E.
Renteria A. 24184—Colombia, Choco, Emisoria La Sirena,
3 km W of La Mansa at top of Cordillera Occidental,
disturbed oak forest, flowers white, elev. 2,300–2,400 m
(16 Jan 1979), (COL!); P.G. Huertas and P.L. Camargo
6679—Colombia, Cundinamarca, Junı́n, Vereda La Cumbre, a orillas de la carretera, bosques, elev. 2,700 m (18 Jan
1967), (COL!); J. Cuatrecasas, R.E. Schultes and E. Smith
12754—Colombia, Norte de Santander, Páramo de Tamá,
Vertiente de Samaria, elev. 2,600–2,900 m (29 Oct 1941),
(COL!).
Notes: The species appears to be related to Gomphichis
costaricensis, from which it is easily separable by the
petals and lip form. The petals of G. lozanoi are cuneate at
the base (vs. sessile, with broad base in G. costaricensis).
The lip of the new species is concave in the lower half,
with two cushion-like, papillate thickenings, divided by
prominent furrow, in contrast to G. costaricensis which is
pubescent in the centre, with very large callus, sulcate in
front.
Eight new species of Gomphichis
Discussion
Spiranthoid orchids comprise a wide spectrum of species
somehow complex for taxonomists to identify. With few
exceptions, all of them are most difficult for proper identification at the species level. Most of them are indistinguishable using solely vegetative characters. In most cases
they do not contain unique or peculiar information sufficient to ensure proper identification of the plant. This can
be seen in a previous section: neither of the vegetative
features can be used to discriminate at the species level in
Gomphichis. Even with the flowers it is impossible to
arrive at an appropriate name without dissecting the usually
tubular flowers. Our experiences in working with spiranthoid species permit us to call the attention of future
students to some items important in the taxonomy of
Gomphichis.
The shape of the lip and alternately its lobation as well
as the presence of thickenings, outgrowth(s) and surface
cover seem to be constant at the species level. Equally
important are the form, length–width ratio and margin
cover of petals, as well as the deflection and cover of the
gynostemium. The reproductive structure of Gomphichis
was described in detail by Szlachetko and Rutkowski
(2000). Some of the characters aforementioned, however,
are observable under lenses only. Combinations of these
features are the most prognostic and useful in determination of the species of Gomphichis.
Occurrence of species described in this paper is characterized by many disjunctions, which could result from
their relationships with paramo. Another fact is that the
very light seeds of Gomphichis can have a long-distance
wind transport and may arrive at distant localities. Their
area of occurrence could be larger during the last glacial
period, when the subparamo zone extended from ca.
2,000 m, with the result that distribution of the new species
could be more continuous. The present-day subparamo belt
extends from 3,200–3,300 to ca. 3,600 m, directly above
the upper forest line (Hooghiemstra et al. 2006).
Paramos are a megadiverse biome with many endemic species. As a proper biome paramos exist since 2–4
millions of years. They have experienced immigrations
from lower altitudes (savanna, upper montane rain forests) and from southern (puna, Austral–Antarctic region)
and northern latitudes via Central America (Hooghiemstra et al. 2006). All this has resulted in the world’s
most megadiverse equatorial tropicalpine biome. Since
the recent past of the Pliocene, about 40 glacial periods
alternating with warmer interstadials have been strongly
operational in speciation processes of the paramo flora
(Hoorn et al. 2010; Van der Hammen and Cleef 1986;
Torres et al. 2013; Smith and Cleef 1988; Cleef 1979).
The paramo reflects a sort of island archipelago, isolated
and fragmented, which promotes high speciation and n
high endemism at the species and genus level (Murillo
1951; Luteyn 1999; Sklenář and Ramsay 2001). Most
endemic species are the result of the repeated island-like
distribution in the Pleistocene and connectivity again
during glacial times (Van der Hammen and Cleef 1986).
Hughes and Eastwood (2006) suggested that rates of
diversification found for high Andean plants driven by
ecological opportunity in island-like model may indeed
outpace the mainland type. Referring to the novelties in
Gomphichis presented in this paper, there is a close
relationship with insularity and altitudinal sequence of
montane forest and paramos.
According to the IUCN Red List (IUCN 2011) for all
species described in this paper, the conservation status is
proposed. Categories were determined based on geographical range of species showing that four species are in
threatened categories and four are in a ‘NT’ category. In
our opinion, keeping in mind the many factors which
determine the survival of terrestrial orchid group of plants,
which is one of the largest in species number threatened by
extinction (Swarts and Dixon 2009), the designated conservation statuses are justified. The main factors having an
impact upon terrestrial orchid population (or species) are
the presence of mycorrhiza, support of pollinators, ecological stability and climate change (Arditti et al. 1990;
Arditti and Ghani 2000; Swarts and Dixon 2009). Ecological stability and climate change belong to long-term
impacting factors. The effects of the latter can be observed
after 20–50 years (Swarts and Dixon 2009; Cleef 2013).
However, it is possible to draw conclusions by observation
of the plant formations where orchids occur, which are part
of ecological stability. Newly described species occur
mainly in the subparamo and grass paramo in Colombia.
The upper forest line is in general close, and the first evidence of displacement of subparamo vegetation by climatic
warming up already has been observed (Cleef 2013). At
present, paramos suffer from multiple human influences
and even destruction of entire habitats by mining, burning
and grazing by cattle and large-scale potato agriculture.
Even the water resources, the most important paramo
ecosystem service, is also strongly affected (in quality and
quantity).
Acknowledgments The senior author would like to thank Dr.
Carlos Parra, the curator of COL, for his support during the visits to
the herbarium. We wish to thank an anonymous reviewer for the
critical comments. The paper is a part of a project supported by a
grant from the Polish Ministry of Science and Higher Education
(8124/B/PO1/2011/40).
Open Access This article is distributed under the terms of the
Creative Commons Attribution License which permits any use, distribution, and reproduction in any medium, provided the original
author(s) and the source are credited.
123
S. Nowak et al.
References
Álvarez A, Cameron KM (2001) Phylogenetic analysis of Gomphichis
and the subtribe Prescottiinae (Orchidaceae). Botanical Society
of America Annual Meeting, Albuquerque
Álvarez-Molina A, Cameron KM (2009) Molecular phylogenetics of
Prescottiinae s.l. and their close allies (Orchidaceae, Cranichideae) inferred from plastid and nuclear ribosomal DNA
sequences. Amer J Bot 96:1020–1040
Arditti J, Ghani AKA (2000) Numerical and physical properties of
orchid seeds and their biological implications. New Phytol
145:146–569
Arditti J, Ernst R, Wing Yam T, Glabe C (1990) The contribution of
orchid mycorrhizal fungi to seed germination: a speculative
review. Lindleyana 5:249–255
Armenteras D, Gast F, Villareal H (2003) Andean forest fragmentation and the representativeness of protected natural areas in the
eastern Andes, Colombia. Biol Conserv 113(2):245–256
Bernal R, Celis M, Gradstein R (2007) Plant diversity of Colombia
catalogued. Taxon 56:273–274
Cleef AM (1979) The phytogeographical position of the neotropical
vascular paramoflora with special reference to the Colombian
Cordillera Oriental. In: Larsen K, Holm-Nielsen LB (eds)
Tropical botany. Academic Press, London, pp 175–184
Cleef AM (2013) Aporte de la misón de los jardines botánicos de
Colombia frente al cambio climático. In: Vásquez CA (ed)
Memorias II Encuentro internacional de ecologı́a aplicada a la
conservación de la flora. Los jardines botánicos afrontando el
cambio climático. Jardı́n Botánico José Celestino Mutis, Bogotá,
D.C., Colombia, pp 35–48
Davis SD, Heywood VH, Herrera-MacBryde O, Villa-Lobos J,
Hamilton A (1997) Centres of plant diversity: a guide and
strategy for their conservation, vol 3. The Americas. IUCN
Publications Unit, Cambridge, England
Dressler RL (1990) The Spiranthoideae: grade or subfamily?
Lindleyana 5:110–116
Dressler RL (1993) Phylogeny and classification of the orchid family.
Dioscorides, USA
Forero E (1988) Botanical exploration and phytogeography of
Colombia: past, present and future. Taxon 37:561–566
Holmgren PK, Holmgren NH (1998) Index herbariorum. New York
Botanical Garden, New York
Hooghiemstra H, Wijninga VM, Cleef AM (2006) The paleobotanical
record of Colombia: implications for biogeography and biodiversity. Ann Missouri Bot Gard 93(2):297–325
Hoorn C, Wesselingh FP, ter Steege H, Bermudez MA, Mora A,
Sevink J, Sanmartı́n I, Sanchez-Meseguer A, Anderson CL,
Figueiredo JP, Jaramillo C, Riff D, Negri FR, Hooghiemstra H,
Lundberg J, Stadler T, Särkinen T, Antonelli A (2010) Amazonia
through time: Andean uplift, climate change, landscape evolution, and biodiversity. Science 330(6006):927–931
Hughes CE, Eastwood RJ (2006) Island radiation on a continental
scale: exceptional rates of species diversification after uplift of
the Andes. Proc Natl Acad Sci USA 103:10334–10339
123
IUCN (2011) Guidelines for using the IUCN red list categories and
criteria: version 9.0. IUCN Species Survival Commission,
IUCN, UK
Kress NJ (1986) The systematic distribution of vascular epiphytes: an
update. Selbyana 9:2–22
Luteyn JL (1999) Páramos: a checklist of plant diversity, geographical distribution, and botanical literature. New York Botanical
Garden, New York
Miani PF, Fajardo M (2001) The integration of biodiversity into
national environmental assessment procedures, national case
studies. UNEP-BPSP Thematic Studies, Colombia
Mittermeier RA, Robles Gil P, Hoffman M, Pilgrim J, Brooks T,
Mittermeier CG, Lamoreux J, da Fonseca GAB (2004) Hotspots
revisited. CEMEX, Mexico, pp 73–82
Murillo LM (1951) Colombia, un archipiélago biológico. Revista
Acad Colomb Ci Exact 9:36–37
Mytnik-Ejsmont J, Szlachetko DL, Nowak S (2012) New species of
the genus Myrosmodes (Orchidaceae, Spiranthoideae) from
Colombia. Pl Syst Evol 298(10):1909–1916
Ortiz Valdivieso P, Uribe Vélez C (2007) Galerı́a de Orquı́deas de
Colombia (CD edition): Asociación Bogotana de Orquideologı́a,
Bogotá
Porup J, Raub K, Reid R, Soriano C (2009) Colombia. Lonely Planet
5th edition, Oakland, p 352
Pridgeon AM, Cribb PJ, Chase MW, Rasmussen FN (2003) Genera
Orchidacearum volume 3, Orchidoideae (part 2), Vanilloideae.
Oxford University Press, Oxford
Sklenář P, Ramsay PM (2001) Diversity of zonal páramo plant
communities in Ecuador. Divers Distrib 7(3):113–124
Smith JMB, Cleef AM (1988) Composition and origins of the world’s
tropicalpine floras. J Biogeogr 15:631–645
Swarts ND, Dixon KW (2009) Terrestrial orchid conservation in the
age of extinction. Ann Bot 104(3):543–556
Szlachetko DL (1995) Systema Orchidalium. Fragm Flor Geobot
Suppl 3:1–152
Szlachetko DL, Nowak S (2013) Two new species of Aa (Orchidaceae, Spiranthoideae) from Colombia. Pl Syst Evol (in press)
Szlachetko DL, Rutkowski P (2000) Gynostemia Orchidalium.
Volume 1. Apostasiaceae, Cypripediaceae, Orchidaceae (Thelymitroideae to Vanilloideae). Acta Bot Fen 169:234
Szlachetko DL, Nowak S, Paluchowska K (2013) Three new species
of the genus Gomphichis (Orchidaceae, Spiranthoideae) from
Colombia. Ann Bot Fenn 50(5):317–323
Torres V, Hooghiemstra H, Lourens L, Tzedakis PC (2013) Astronomical tuning of long pollen records reveals the dynamic
history of montane biomes and lake levels in the tropical high
Andes during the quaternary. Quatern Sci Rev 63:59–72
Van der Hammen T, Cleef AM (1986) Development of the high
Andean páramo flora and vegetation. In: Vuillemuier F, Monasterio M (eds) High altitude tropical biogeography. Oxford
University Press, Oxford, pp 153–200
Joanna Mytnik