Eight new species of Gomphichis (Orchidaceae, Spiranthoideae) from Colombia

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.


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 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 . 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, 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 (Á lvarez and Cameron 2001;Á lvarez-Molina and Cameron 2009;Dressler 1990Dressler , 1993Szlachetko 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 (Mytnik-Ejsmont et al. 2012;. The collected specimens are identified based on the comparison with the type material and original descriptions. The results of our recent herbarium studies (2009)(2010)(2011)(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, and Kolanowska (2009Kolanowska ( -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 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
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.
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.
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.  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.
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, 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 ovateelliptic, 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.
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 Eight new species of Gomphichis 123 vulnerable (VU B1ab(i,iii)). The species is known only from five localities. An extent of its occurrence is about 39,000 km 2 , 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 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 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) 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). 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.
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 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).
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 km 2 and the known localities are all restricted to very narrow elevation range in high Andean paramo.
Representative 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.
Etymology: Dedicated to Roberto Jaramillo Mejía, 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 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.
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.
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.
Ecology: Terrestrial in oak forest, also in páramo. Flowering time: August, October-January.
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 km 2 , 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 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.

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).