Introduction

The list of Colombian orchids compiled by Ortiz Valdivieso and Uribe Vélez (2007) includes almost 3,000 species and subspecies. This number seems to be underestimated taking into consideration the latest discoveries of new taxa representing various taxonomic groups (Higgins and Viveros 2008; Giraldo and Dalström 2012; Hills and Weber 2012; Kolanowska and Pérez-Escobar 2012; Kolanowska and Szlachetko 2012, 2013a, b; Kolanowska et al. 2012; Pérez-Escobar et al. 2013; Szlachetko and Baranow 2012; Szlachetko et al. 2013). The best illustration of this situation is the recent revision of Microchilus (Goodyerinae) which resulted in the description of over 90 new species (Ormerod 2002, 2005, 2007, 2008, 2009a, b). The limited monitoring of the tropical regions resulting in the insufficient herbarium representation together with the difficulties related to the small size of the perianth parts, complicated lip construction and hiding of taxonomically important characters inside usually tubular flowers makes many spiranthoid orchids intractable, but promising object of study.

One of the taxa of which specific diversity remains poorly recognized is Cranichis described in the end of XVIII century (Swartz 1788). The nominal species, C. muscosa Sw., was selected over 150 years after the formal description (Acuña 1939). The taxonomic position of the genus in Orchidaceae is not questioned. While in the older classification systems (Pfitzer 1889; Bentham and Hooker 1862–1863; Dressler and Dodson 1960), Cranichis was placed in a broadly defined tribe Neottieae, currently it is recognized as a member of subtribe Cranichidinae in Cranichideae (Dressler 1981, 1993; Szlachetko 1995; Szlachetko and Rutkowski 2000; Chase et al. 2003). Nevertheless, Dressler (1981, 1993) and Szlachetko (1995) considered Cranichideae as a tribe of subfamily Spiranthoideae based on their morphological characters, while the molecular taxonomists (Chase et al. 2003; Salazar et al. 2003) placed it in Orchidoideae.

The aim of this study was to evaluate the morphological differences between Cranichis and allied genera segregated from this taxon in the recent years: Exalaria Garay and Romero, Ocampoa A. Rich. and Gal. and Pseudocranichis Garay. In addition, the results of the revision of Colombian representatives of Cranichis s.s. are presented and a key to the national species is provided.

Materials and methods

In total about 1,000 herbarium specimens of Cranichis and related genera were revised. The material was studied in or borrowed from the following institutions: AAU, AMES, AMO, B, BM, C, CAS, CAY, CICY, COL, CUVC, E, ENCB, F, FLAS, G, GB, HAJB, HB, HEID, HPUJ, HUA, HURP, INB, JAUM, K, L, LE, LINN, M, MA, MEXU, MO, MOL, NY, P, QCA, QCNE, QPLS, RENZ, RPSC, S, SEL, TEFH, TULV, U, UC, UGDA, UPRRP, US, USJ, USM, VALLE, VEN and W.

Dried herbarium specimens were examined according to the standard procedures. Every studied sheet was photographed and the data from the labels were taken. Both vegetative and generative characters of each plant were studied. The shape and size of the leaves were examined first, then the arrangement of the inflorescence and the shape and size of the floral bracts and ovary. The morphology of flower and the surface of its elements were examined after softened in boiling water. The stereomicroscope was used to make observations of the perianth parts and their measurements.

Acronyms for herbaria cited in this paper followed Index Herbariorum (Thiers continuously updated). The CorelDraw v.12 software was used for the preparation of the distribution maps.

Classification of cranichid orchids

Cranichidinae sensu Dressler (1981) comprises 15 genera grouped in two alliances corresponding more or less with his later subtribes Prescottinae and Cranichidinae s.str. (Dressler 1993). Both subtribes differ considerably one from another in the gynostemium and pollinia structure. Prescottinae are characterized by large, bi-winged clinandrium, completely covering the sides and back of the anther, or occasionally vestigial. Stigma is entire to separately bilobed, with lateral lobes well developed to rudimentary and rostellum is shelf-like, soft, thin, erect, obtuse or truncate at the apex. Pollinia in orchids of this group are soft and powdery. Cranichidinae s.str. can be determined as having clinandrium usually stretched between the filament and the apical margin of the stigma, usually detached from the anther. Stigma is horizontal, transversely elliptic to gibbous, deeply concave, and formed of the lateral lobes of the stigma. The prominent rostellum is usually digitate, fleshy, erect, and acute. Pollinia are relatively hard, formed of compact pollen grains.

In both groups the flowers are non-resupinate, but in Prescottinae the lip is usually concave, often fringed along the margins and in Cranichidinae it is fixed, saccate to spurred basally.

Based primarily on the gynostemium structure, mainly rostellum, viscidium and pollinium organization Szlachetko (1995) proposed different placement of considered subtribes. He included Prescottinae to Spirantheae, leaving Cranichideae as a monotypic tribe. Salazar et al. (2009) based on analyses of selected DNA markers combined genera of both subtribes into Cranichidinae s.l.

The subtribe Prescottinae Dressl

Aa Rchb.f., Altensteinia Kunth, Galeoglossum A. Rich. and Gal., Gomphichis Lindl., Myrosmodes Rchb.f., Porphyrostachys Rchb.f., Prescottia Lindl., Pseudocranichis Garay, Stenoptera C. Presl

The subtribe Cranichidinae Lindl

Baskervilla Lindl., Cranichis Sw., Exalaria Garay and Romero, Fuertesiella Schltr., Ocampoa A. Rich. and Gal., Ponthieva R.Br., Pseudocentrum Lindl., Pterichis Lindl., Solenocentrum Schltr.

Taxonomic concepts of Exalaria, Ocampoa and Pseudocranichis

While most of the species included in Cranichis share similar vegetative and floral characters (Figs. 1, 2a, 3a, b), the taxonomic affinities of the three of them, e.g. C. thysanochila Rob. and Greenm., C. mexicana (A. Rich. and Galeotti) Schltr. and C. fertilis (F. Lehm. and Kraenzl.) Schltr., have been recently discussed and none of these taxa is treated as Cranichis at the present day. Their generic affinities are here discussed.

Fig. 1
figure 1

Cranichis engelii. a Dorsal sepal, b petal, c lateral sepal, d lip, and e lip (side view). Drawn by A. Król

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Fig. 2
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Gynostemium morphology of Cranichidinae representatives: Cranichis (a), Exalaria (b, c), Ocampoa (d–f), Ponthieva (g, h) and Baskervilla (I)

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Fig. 3
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a, b Cranichis ciliata (Photos by A. Kay); c, d Exalaria parviflora (Photos by M. Kolanowska and A. Kay); e, f Ocampoa mexicana (Photo by C. Velazco); g, h Pseudocranichis thysanochila Holotype; i, j Galeoglossum tubulosum (Photos by F. Muller); k, l. Ponthieva racemosa (Photos by A. Kay)

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The first one, Mexican C. thysanochila (Fig. 3g, h) is distinguished by the bipartite lip with a fringed apex, two thickened and pubescent veins (Fig. 4) on the disc and gynostemium (Fig. 5a–c) with saddle-shaped stigma and short, truncate rostellum. As this species morphologically does not correspond to any other cranichid genus it was elevated to the generic rank by Garay (1982) and named Pseudocranichis. The gynostemium morphology (Szlachetko 1995) and the results of the molecular research (Álvarez-Molina and Cameron 2009) confirmed the close relation of Pseudocranichis thysanochila with Prescottiinae rather than Cranichidinae. In the genetic level, this species is nested in Prescottia, with strong support for its position as sister to P. tubulosa (Figs. 6, 5d, e, 3i, j) from which it differs significantly in the morphological characters. Salazar (2009) proposed the transfer of P. thysanochila to the recently restituted genus Galeoglossum A. Rich. and Galeotti, together with Prescottia tubulosa (Lindl.) L.O. Williams, considering this taxon as a synonymic with Galeoglossum prescottioides A. Rich. and Galeotti. Worth to mention that Prescottia tubulosa was described initially also as a member of Cranichis. Both P. thysanochila and Galeoglossum tubulosum are restricted in their distribution to the conifer–oak forests in the mountains of Mexico and Guatemala and they may grow sympatrically (Salazar 2009). Both, according to Salazar (2009), are distinguished from other cranichid orchids by a saddle-shaped stigma with a wet, sticky receptive area at each side, being separated by a dry, non-receptive central portion. Other characters shared by those species are tepals recurved above the middle, the lip base provided with a retrorse, rounded auricle on each side and the ribbon-like pollinia joined at the apex to one another and to a small, ovate or deltate viscidium. Both species, however, differ significantly in the flower morphology. In Pseudocranichis tepals are free to the base and the lip is distinctly constricted in the apical third or quarter with a prominent, lacerate apical lobule (Fig. 4). The tepals of Galeoglossum are basally connate and lip apical lobule is inconspicuous and entire (Fig. 6). Considering these differences, the generic separation of Pseudocranichis from Galeoglossum seems to be justified.

Fig. 4
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Pseudocranichis thysanochila. a Dorsal sepal, b petal, c lateral sepal, d lip, and e flower (side view). Drawn by D.L. Szlachetko

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Fig. 5
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Gynostemium morphology of Prescottinae representatives: Pseudocranichis (a–c), Galeoglossum (d, e) and Prescottia (f–h)

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Fig. 6
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Galeoglossum tubulosum. a Flower (side view), b dorsal sepal, c petal, d lateral sepals and e lip. Drawn by A. Król

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There is another species which was considered as a member of Cranichis, Ocampoa mexicana A. Rich and Galeotti (Fig. 3e, f). It was embedded into Cranichis by Schlechter (1918); however, this replacement was argued by González Tamayo (1995) and Soto (2008). Ocampoa mexicana differs from Cranichis species by the long-clawed, navicular lip lacking branching, coloured veins, the basally auriculate sepals (Fig. 7) and the gynostemium lacking the wing-like margins (Fig. 2d–f). Its vegetative characters, e.g. concolor, short-petiolate leaves with a conspicuous net of slightly protruding nerves suggest the close relation with Baskervilla Lindl. and Ponthieva R. Br., however, Bentham and Hooker (1862–1863) noticed the similarity of Ocampoa to Prescottia. The form of perianth segments, e.g. asymmetric lateral sepals and linear petals, resembles that of Baskervilla, but from the latter O. mexicana differs by the petals and lip being free from the gynostemium. Ocampoa was usually accepted by the orchidologists (Hágsater et al. 2005; Soto 2008); however, recently Salazar et al. (2009) based on the results of the genetic research proposed a transfer of the species into Ponthieva, despite essential differences in the flower and gynostemium structures between taxa in question. The gynostemium of Ponthieva is stalked, swollen above narrow base, connate with the lip and petals. The petals are strongly asymmetric, clawed and lip variously transformed with various calli.

Fig. 7
figure 7

Ocampoa mexicana. a Flower (side view), b dorsal sepal, c petal, d lateral sepal, e lip, f lip (side view) and g lip and gynostemium in the natural position. Drawn by P. Baranow

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The last species which does not fit the morphological concept of Cranichis is C. fertilis (Figs. 3c, d, 8) widely distributed in tropical South America. The most distinctive characters of the plants include the triangular stigma, the bifid rostellum and the lack of hamulus (Fig. 2b, c, Szlachetko and Rutkowski 2000). Based on those features the species was elevated to the generic rank as Exalaria parviflora by Garay and Romero-González (1999) and this position was confirmed in the molecular studies of Álvarez-Molina and Cameron (2009, Fig. 9). The concept of monotypic Exalaria was argued by Salazar who transferred E. parviflora to Ponthieva (Salazar et al. 2009), despite inconsistency of this replacement with the morphological diagnostic characters of the latter genus which includes adnation of the petals above the gynostemium base and the presence of the clawed lip that is often much smaller than tepals, Moreover, the rostellum of Ponthieva is truncate (Fig. 2g, h), not bifid like in E. parviflora. We believe that since this species cannot be placed within any existing genus, it should be maintain as the monotypic, separate one.

Fig. 8
figure 8

Exalaria parviflora. a Dorsal sepal, b petal, c lateral sepal, d lip and e lip (side view). Drawn by A. Król

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Fig. 9
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Fragment of strict consensus of most parsimonious trees presented by Álvarez-Molina and Cameron (2009)

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As mentioned before both O. mexicana and E. parviflora were reassigned recently by Salazar et al. (2009) to the genus Ponthieva (Figs. 2g, h, 3k, l, 10) based on the results of the molecular studies in which both taxa were placed in the clade that includes the type of Ponthieva. However, the “Ponthieva clade” seems to be divided into two groups—within one of them can be found also Baskervilla colombiana Garay which, obviously, was not considered by the authors as a member of Ponthieva from which it clearly differs by the elongate, slender gynostemium (Szlachetko and Rutkowski 2000). Moreover, while in the Salazar’s strict consensus tree reconstructed by maximum parsimony analysis Ocampoa is sister to Ponthieva ephippium Rchb. f., in the Bayesian summary cladogram O. mexicana is sister to E. parviflora receiving high posterior probability value. The inconsistency between the phylogenetic trees together with the obvious morphological differences between Ponthieva species and considered taxa encourages to maintain them as separate genera. The comparison of the vegetative and floral characters of Cranichis, Exalaria, Ocampoa, Pseudocranichis, Galeoglossum and Ponthieva is presented in Table 1. The variation in the gynostemium morphology between all genera mentioned before is illustrated in Figs. 2 and 5. The photographs of the discussed taxa are presented in Fig. 3.

Fig. 10
figure 10

Ponthieva diptera. a Dorsal sepal, b petal, c lateral sepal, d lip, e lip (side view). Drawn by A. Król

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Table 1 Comparative morphology of Cranichis, Exalaria, Ocampoa Pseudocranichis, Galeoglossum and Ponthieva
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From the genera discussed above until now only Cranichis and Exalaria were reported from Colombia; however, our recent studies revealed the existence of Ocampoa species in the national flora. The results of the recent revision of the Colombian Cranichis with the key to the identification of national representatives of the genus are presented in this paper together with the proposals of the new combination within Ocampoa and Pseudocranichis.

Taxonomic treatment

Considering the usually accepted exclusion of C. thysanochila, C. mexicana and C. fertilis from Cranichis, the genus is clearly defined morphologically. Its representatives are easily distinguished from other cranichid orchids by the villous-hairy roots, the distinctly petiolate, suberect or arcuately spreading leaves, non-resupinate flowers, petals much narrower than sepals, and cochleate lip, often with conspicuously marked, coloured reticulate veins. The gynostemium of those plants is relatively massive, often swollen at the apex, without column-foot. The motile anther is oblong to ovate, 2-chambered and the inconspicuous caudiculae are formed from the apices of pollinia. The clinandrium is usually thick, massive, spacious. The single viscidium is thick and relatively small and the hamulus is usually elongate, finger-like, thick, directed towards the anther (Szlachetko and Rutkowski 2000).

Cranichis representatives grow as terrestrial or lithophytic plants in lowlands as well as in montane forest, usually in humus and Sphagnum tussocks. Some populations were reported as subepiphytic. The altitudinal range of the genus extends from 350 up to 3,000 m a.s.l. (Carnevali and Ramírez-Morillo 2003; Cribb 2003). Currently about 60 known species of the genus are distributed from Florida and Mexico to Bolivia and Argentina. In the Colombian orchid flora 19 species of the genus were found so far. Most of them, except C. muscosa and C. parvula, were reported from the submontane and montane regions of the country (Ortiz Valdivieso and Uribe Vélez 2007). Plants are common in the litter of the Andean forest and heath and shrubs of subparamo and paramo where they grow in damp and mossy areas. The national species were also reported from pastures and rocky slopes (Dueñas Gómez and Fernández Alonso 2009). The last comprehensive revision of Colombian Cranichis was conducted over 50 years ago (Schneider 1953). Despite the wide geographical range, the novelties within Cranichis are rather rare, the most recent description of new species comes from 2004 (Christenson 2004). Our research on Colombian representatives of the genus resulted in description of new entities (Szlachetko and Kolanowska 2013) from this country. We believe that the actual species diversity of Cranichis is underestimated and the studies on those small-flowered, terrestrial orchids should be intensified.

Key to Colombian species of Cranichis

1. Petals with long hairs or ciliae along the margins…….. C. wageneri (1)

1*. Petals without long hairs on the margins……. 2

2. Petals ciliate….. 3

2* Petals with entire or slightly erose margins, glabrous…… 12

3. Scape glabrous……… 4

3* Scape glandular or glandular-pubescent……….. 6

4. Petals from a short claw broadly oblanceolate to elliptic…. C. polyantha (2)

4* Petals linear-ligulate to oblong-oblanceolate……. 5

5. Lip midvein unbranched……. C. engelii (3)

5* All lip veins branching……. C. muscosa (4)

6. Ovary glandular or papillate……. 7

6* Ovary glabrous……. 9

7. Lip with spiculate pustules or knob-like projections……. C. schlimii (5)

7* Lip without any additional projections on the surface……. 8

8. Lip broadly ovate, obtuse, nerves 3, main nerves thickened, branches and anastomozes thin ………. C. brachyblephara (6)

8* Lip elliptic-ovate in outline, apex truncate with short, triangular, acute apiculus ……. C. zarucchii (7)

9. Inflorescence elongate, raceme 8–12 cm long, longer than the leaf blade……. C. antioquiensis (8)

9* Raceme up to 4–7 cm long, shorter than the leaf blade……. 10

10. Ovary about 5–6 mm long……. 11

10* Ovary up to 11 mm long……. C. ciliata (9)

11. Sepals attenuate towards the apex, petals minutely ciliate along margins, lip longer than wide……. C. pleioneura (10)

11* Sepals rounded at the apex, petals pubescent along margins, lip as long as wide……. C. polyblephara (11)

12. Petals margins erose … C. picta (12)

12* Petals margins entire … 13

13. Leaves narrowly oblanceolate to linear-lanceolate … C. lehmanniana (13)

13* Leaves ovate, ovate-lanceolate to orbicular … 14

14. Leaves usually two, often one of them is much larger than another … C. diphylla (14)

14* Leaves several, subequal in size … 15

15. Petals elliptic, widest near the middle … C. gibbosa (15)

15* Petals lanceolate to oblong-oblanceolate, widest near the base or near the apex … 16

16. Floral bracts subequal or longer than ovary … C. lehmannii (16)

16* Floral bracts shorter than ovary … 17

17. Leaf blade elliptic to orbicular; petiole 0.4 cm long … C. tenuis (17)

17* Leaf blade broadly ovate to broadly elliptic; petiole at least 1 cm long … 18

18. Petiole up to 2.5 cm long, lip unlobed … C. parvula (18)

18* Petiole up to 9 cm long, lip 3-lobed at apex … C. pulvinifera (19)

1. Cranichis wageneri Rchb.f.

Linnaea 41: 19. 1877; Type (hoc loco designatus): Venezuela. Wagener s.n. (Lectotype: W!). - Ames and Correll, Fieldiana, Bot. 26(1): 86–87. 1952. - Schweinfurth, Fieldiana, Bot. 30(1): 116-117. 1958. - Foldats in Lasser, Fl. Venezuela, Orchid. 15(1): 422. 1969.

Reference material—Colombia. Valle del Cauca. Mpio. Yumbo. Dapa, Jorge Negret farm, 35 min from el Rodadero station. Alt. 2,074 m. 3°32′54.9″N 76°35′15.4′′W. Apr 2012. E. Parra s.n. (VALLE!).

2. Cranichis polyantha Schltr.

Repert. Spec. Nov. Regni Veg., Beih. 7: 61. 1920; Type (Garay 1978: 203): Colombia. Madero 22 (B†, Lectotype: AMES).

Reference material—Colombia. Cauca. Sine loc. Alt. 2000 m. Madero s.n. (B†); Sine loc., Madero 22 (B†, AMES).

3. Cranichis engelii Rchb.f.

Linnaea 41: 19. 1876; Type (Garay 1978: 196): Venezuela. Merida. Engel s.n. (Lectotype: W!).

Reference material—Colombia. Norte de Santander. Sine loc. (Schlechter 1920).

4. Cranichis muscosa Sw.

Prodr.: 120. 1788; Type (Garay 1978: 202): Jamaica. Swartz s.n. (Lectotype: BM!; Isotype: W!).

Reference material—The occurrence of this species in Colombia was reported by Dueñas Gómez and Fernández Alonso (2009).

5. Cranichis brachyblephara Schltr.

Repert. Spec. Nov. Regni Veg., Beih. 7: 58. 1920; Type (hoc loco designatus): Colombia. Madero s.n. (B†).

Reference material—Colombia. Cauca. Sine loc. Alt. 3,000 m. Madero s.n. (B†).

6. Cranichis zarucchii Szlach. and Kolan.

Plant Syst. Evol. 299: 981. 2013; Type: Colombia. Zarucchi, Brant and J. Castano 5694 (Holotype: COL!).

Reference material—Colombia. Antioquia. Mpio. Sonsón. Km 11 of road Sonsón-Nariño. 25 km from Narino, near km post 151 marking distance from Bogotá. Pluvial subparamo vegetation. 542′N, 7515′W. Alt. 2,780 m. 1 Apr 1987. Zarucchi, Betancur and F.J. Roldan 5177 (COL!).

7. Cranichis schlimii Rchb.f.

Linnaea 41: 19. 1877; Type (Garay 1978: 206): Venezuela. Funck and Schlim 1238 (Lectotype: W!).

Reference material—Colombia. Valle del Cauca. Mpio. El Cerrito. I.P. Auji, hacienda La Pajosa. Alt. 2,800 m. 22 May 1990. S. Sarria, M.T. Sierra and J. Rivas 1422 (CUVC!).

8. Cranichis antioquiensis Schltr.

Repert. Spec. Nov. Regni Veg., Beih. 7: 57. 1920; Type (Garay 1978: 189): Colombia. Madero 168 (B†; Lectotype: AMES-drawing).

Reference material—Colombia. Antioquia. Sine loc. Alt. 2,000 m. Madero s.n. (B†); Sine loc. Madero 168 (B†; AMES-drawing).

9. Cranichis atrata Schltr.

Repert. Spec. Nov. Regni Veg., Beih. 7: 58. 1920; Type: Colombia. Madero s.n. (B†).

Reference material—Colombia. Cauca. Sine loc. Madero s.n. (B†).

10. Cranichis pleioneura Schltr.

Repert. Spec. Nov. Regni Veg., Beih. 7: 60. 1920; Type: Colombia. Madero s.n. (B†).

Reference material—Colombia. Cauca. Sine loc. Alt. 2,500 m. Madero s.n. (B†).

11. Cranichis polyblephara Schltr.

Repert. Spec. Nov. Regni Veg., Beih. 7: 61. 1920; Type: Colombia. Lehmann 2812 (B†).

Reference material—Colombia. Cauca. Bei Popayán. Alt. 1,600–1,900 m. May 1883. Lehmann 2812. (B†).

12. Cranichis picta Rchb.f.

Linnaea 41: 52. 1877; Type (Garay 1978: 203): Ecuador. Jameson s.n. (Lectotype: W!).

Reference material—Colombia. Volcan de Pasto. Alt. 3,000 m. 14 Feb 1880. Lehmann 476 (W!).

13. Cranichis lehmanniana (Kraenzl.) L.O. Williams

Lilloa 3: 477. 1938. - Goodyera lehmanniana Kraenzl., Bot. Jahrb. Syst. 26: 498. 1899; Type (Garay 1978: 199): Colombia. Lehmann 6149 (Lectotype: K!).

Reference material—Colombia. Nariño. Lehmann 6149 (K!). Cauca. Sine loc. (Schlechter 1920).

14. Cranichis diphylla Sw.

Nov. Gen. Sp. Prodr.: 120. 1788; Type (Garay 1978: 192): Jamaica. Swartz s.n. (Lectotype: BM!; Isotype: W!).

Reference material—The occurrence of this species in Colombia was reported by Dueñas Gómez and Fernández Alonso (2009).

15. Cranichis gibbosa Lindl.

Ann. Nat. Hist. 15: 385. 1845; Type (Garay 1978: 198): Ecuador. Hartweg 1435 (Lectotype: K-L!, Isotypes: AMES, K!, NY, W!).

Reference material—the occurrence of this species in Colombia was reported by Ortiz Valdivieso and Uribe Vélez (2007).

16. Cranichis lehmannii Rchb.f.

Otia Bot. Hamb. 1: 4. 1878; Type (Garay 1978: 199): Ecuador. Lehmann 77 (Lectotype: W!).

Reference material—Colombia. Antioquia. Mpio. Frontino, Corregimiento de Nutibara, zona de Murrí, Alto de Cuevas. Alt. 1,000–1,850 m. 14 Feb 1991, R. Callejas and al. 9899 (HUA—Dueñas Gómez and Fernández Alonso 2009).

17. Cranichis tenuis Rchb.f.

Flora 48: 274. 1865; Type (hoc loco designatus): Cuba. Wright 1478 (Lectotype: W!).

Reference material—The occurrence of this species in Colombia was reported by Ortiz Valdivieso and Uribe Vélez (2007).

18. Cranichis parvula Renz

Candollea 11: 259, Fig. 8b. 1948; Type (Garay 1978: 202): Colombia. Renz 4159 (RENZ).

Reference material—Colombia. Boyacá. Alt. 500 m. 10 Oct 1938. Renz 4159 (RENZ). Meta. Alt. 500–700 m. 10 Oct 1939. Renz 4124 (RENZ).

19. Cranichis pulvinifera Garay

Fl. Ecuador 9: 204, Fig. 61 A. 1978; Type: Colombia. Bristol 1227 (Holotype: AMES).

Reference material—Colombia. Putumayo. Bristol 1227 (AMES).

Taxonomic transfers

Our study on cranichid orchids revealed the necessity of three taxonomic transfers.

Pseudocranichis cactorum (Salazar and Chávez-Rendón) Szlach. and Kolan., comb. Nov.

Basionym: Galeoglossum cactorum Salazar and Chávez-Rendón, Syst. Bot. 36(2): 262. 2011.

Notes. The lateral sepals of this species are free and its lip is distinctly constricted in the apical part with a prominent, dentate apical lobule. The close relationship of this species with P. thysanochila was also confirmed in the molecular tree presented by Salazar et al. (2011).

Ocampoa carlos-parrae (Szlach. and Kolan.) Szlach. and Kolan., comb. Nov.

Basionym: Cranichis carlos-parrae Szlach. and Kolan., Pl. Syst. Evol. 299: 979. 2013.

Ocampoa crumenifera (Garay) Szlach. and Kolan., comb. Nov.

Basionym: Cranichis crumenifera Garay, Caldasia 8: 518. 1962.

Notes. Both Cranichis carlos-parrae and C. crumenifera should be transferred to Ocampoa based on the presence of C-shaped, long lip claw and basally auriculate sepals.