Cranichis badia, C. brevirostris and C. silvicola spp. nov. (Orchidaceae, Cranichidinae) from Colombia and Venezuela

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Nordic Journal of Botany 000: 001 009, 2013 doi: 10.1111/j.1756-1051.2013.00172.x 2013 The Authors. Nordic Journal of Botany 2013 Nordic Society Oikos Subject Editor: Bertil Ståhl. Accepted 4 March 2013 Cranichis badia, C. brevirostris and C. silvicola spp. nov. (Orchidaceae, Cranichidinae) from Colombia and Venezuela Marta Kolanowska and Dariusz L. Szlachetko M. Kolanowska (martakolanowska@wp.pl) and D. L. Szlachetko, Dept of Plant Taxonomy and Nature Conservation, Univ. of Gdańsk, ul. Wita Stwosza 59, PL-80-308 Gdańsk, Poland. Three new species of the genus Cranichis first recognized by Jany Renz but never published, C. badia, C. brevirostris and C. silvicola, are described and illustrated. Information about their ecology and distribution is provided together with a brief discussion on their related species. The genus Cranichis was described by Olof Swartz (1788) based on the characteristic cochleate lip positioned uppermost in the non-resupinate flower, thus, resembling a helmet (Greek: kranos). As there was no obligation to designate a generitype until 1958, the nominal species was selected, viz. C. muscosa, over 150 years after the genus was described (Acuña 1939). Despite the problematic relationships among spiranthoid orchids, especially among Cranichidinae and Prescottiinae (Dressler 1981, 1993, Szlachetko 1995, Chase et al. 2003) and disagreement over delimitation of genera within those taxa, the distinctness of Cranichis is unquestionable considering both morphological (Dressler 1993, Szlachetko and Rutkowski 2000) and molecular (Álvarez-Molina and Cameron 2009) studies. Species of this genus are easily distinguished from other Cranichidinae by the villoushairy roots, the distinctly petiolate, suberect or arcuately spreading leaves, non-resupinate flowers, petals much narrower than sepals, and the fleshy, cochleate lip, often with conspicuous, coloured, reticulate veins. Since the transfer of C. fertilis (F. Lehm. & Kraenzl.) Schltr. to Exalaria Garay & G. A. Romero-Gonzalez (Garay and Romero-González 1999), the genus may also be clearly defined by its gynostemium morphology. This structure is relatively massive, often swollen at the apex and it is lacking a column-foot. The motile anther is oblong to ovate, 2-chambered. The inconspicuous caudiculae are formed from the apices of pollinia. The clinandrium is usually thick, massive and spacious. The single viscidium is relatively small and thick, the hamulus is usually elongate, finger-like, thick, and directed towards the anther (Szlachetko and Rutkowski 2000). Cranichis is currently comprised of ca 60 species, distributed from Florida and Mexico to Bolivia and Argentina. The wide geographical range reflects the ecological variation of those plants which grow as terrestrials or lithophytes in lowlands as well as in montane regions. They are usually found inside forests, commonly in humus and Sphagnum tussocks. The altitudinal range of the genus extends from 350 m up to 3000 m a.s.l. (Carnevali and Ramírez-Morillo 2003, Cribb 2003). Despite the wide geographical range, novelties within Cranichis are rather rare. The most recent description of a new species was done in 2004 (Christenson 2004). Limited monitoring of the tropical regions results in insufficient herbarium collections. This fact, together with problems in precise examination of dried Cranichis plants due to their small-sized flowers and complicated (cochleate) lip construction, makes Cranichis difficult for taxonomic studies. During studies on north-andean orchids we came across material of three distinctive Cranichis species which were preliminary recognized by Jany Renz who placed short notes on their sheets deposited in RENZ. Since he never effectively published his findings (Art. 29.1 ICBN, McNeill et al. 2006), the complete descriptions, illustrations and notes on the taxonomic affinities of the new species are presented in this paper. Cranichis badia Renz ex Kolan. & Szlach. sp. nov. (Fig. 1) Most similar to C. engelii Rchb. f. from Ecuador, Colombia and Venezuela, but separable from it by ligulate-lanceolate to lanceolate petals (vs petals obliquely oblong-oblanceolate to linear-oblanceolate), and elliptic-subrhombic, subacute to obtuse lip (vs lip ovate on prominent clawed-like lower part, rounded or truncate at the apex). Type: Venezuela, Est. Mérida, 2300 m a.s.l., 7 Nov 1949, O. Renz 6065 (holotype: RENZ!, isotypes: RENZ!). Early View (EV): 1-EV see http://tinyurl.com/ctujsze

Figure 1.Cranichis badia. (A) dorsal sepal, (B) petal, (C) lateral sepal, (D) lip, (E) lip, side view. Scale bar 2 mm. Drawn by A. Król from the holotype. Description Plants up to 40 cm tall, erect. Leaves 1 3, gathered in a basal rosette, petiolate; petiole 5 15 cm long, narrow, canaliculate; blade 4 12 cm long and 2 5 cm wide, elliptic to ovateelliptic, acuminate, cuneate at base. Scape up to 32 cm long; rachis up to 15 cm long, subdensely many-flowered. Flowers small, brown; lip white with brown venation. Floral bracts 5 6 mm long, lanceolate, acuminate. Pedicellate ovary 8 10 mm long. Sepals 3-nerved. Dorsal sepal 4.0 4.5 mm long, 1.8 mm wide, elliptic to oblong-ovate, subacute. Lateral sepals 3.5 mm long, 2.8 mm wide, obliquely oblong-elliptic to elliptic, subobtuse to subacute. Petals 3.2 4.0 mm long, 0.4(1.2) mm wide, linear, almost filiform, rounded at apex and with glabrous margins. Lip up to about 3 mm long, 2.5 3.0 mm wide, cochleate, elliptic-subrhombic, subacute to obtuse at apex, with the midvein anastomosing into strongly branching lateral veins. Gynostemium 1.6 2.0 mm long. Ecology Cranichis badia grows at about 2300 2900 m a.s.l. Flowering in November and December. Distribution Cranichis badia is known from Colombia and Venezuela (Fig. 7). Similar species Cranichis badia is similar to C. engelii Rchb. f. known from Ecuador, Colombia and Venezuela, but differs mainly by the ligulate-lanceolate to lanceolate petals (vs petals obliquely oblong-oblanceolate to linear-oblanceolate), and 2-EV

Figure 2. Comparison of the floral elements of (A) Cranichis badia, Renz 6065, RENZ, (B) C. engelii, Bruckmuller s.n., W, (C) C. atrata, Madero 3, AMES. Drawn by A. Król. 3-EV

Figure 3.Cranichis brevirostris. (A) dorsal sepal, (B) petal, (C) lateral sepal, (D) lip, (E) lip, side view. Scale bar 2 mm. Drawn by A. Król from the holotype. elliptic-subrhombic, subacute to obtuse lip (vs lip suborbicular, rounded or truncate at the apex). Another species similar to C. badia is C. atrata Schltr., but the petals are very narrow, linear, with glabrous margins in the former (vs oblong-lanceolate with long-ciliate margins), and sepals are 3-nerved (vs 1-nerved) (Fig. 2). The new species is easily distinguished from other species with glabrous petals (Cranichis diphylla Sw., C. fendleri Schltr., C. lehmannii Rchb. f., C. muscosa Sw., C. tenuis Rchb. f.) by the brown flowers with clawed, elliptic-subrhombic, subacute to obtuse lip. Additionally, from C. diphylla it differs by the leaves cuneate at the base with elliptic to ovate elliptic blade (vs base cordate, blade ovate to broadly ovate), from C. muscosa by the linear, almost filiform petals (vs petals linear-ligulate to narrowly-oblanceolate) and from C. tenuis by the leaf shape and size (4 12 2 5 cm vs 0.9 6.0 0.6 2.0 cm). Additional specimens examined (paratypes) Colombia, Dept Cundinamarca, Bogotá, Quebrada del Chicó, 2700 2750 m a.s.l., 27 Dec 1943, M. Schneider 172/5 (RENZ). Venezuela, Est. Tachira, 2880 m a.s.l., 23 Nov 1949, O. Renz 6174 (RENZ), the same location, 2900 m a.s.l., 23 Nov 1949, O. Renz 6175 (RENZ). Cranichis brevirostris Renz ex Kolan. & Szlach. sp. nov. (Fig. 3) Easily distinguishable from C. monophylla Lindl. by the crenate margins of the leaves, the suborbicular lip, devoid 4-EV

Figure 4. Comparison of the floral elements of (A) Cranichis brevirostris, Renz 5960, RENZ and (B) C. monophylla, Fendler 2138, K. Drawn by A. Król. 5-EV

Figure 5.Cranichis silvicola. (A) dorsal sepal, (B) petal, (C) lateral sepal, (D) lip, (E) lip, side view. Scale bar 2 mm. Drawn by A. Król from the holotype. of any callosities (vs lip ovate-oblong, adorned with numerous fleshy projections on the inner surface), subdensely-flowered inflorescence (vs rachis laxly flowered) and floral bracts longer than ovary (vs floral bracts shorter than ovary). Type: Venezuela, Est. Tachira, 2700 m a.s.l., 20 Sep 1949, O. Renz 5960 (holotype: RENZ!, isotypes: RENZ!). Description Plants up to 35 cm tall, erect. Leaves 1 2, gathered in a basal rosette, petiolate; petiole 3.75 12.00 cm long, narrow, canaliculate; blade 2.6 11.0 cm long and 3.25 4.75 cm wide, elliptic to ovate, acute or shorty acuminate, cuneate at base, with crenate margins. Scape up to 28 cm long; rachis 2.50 5.75 cm long, subdensely many-flowered. Floral bracts 7 8 mm long, lanceolate, acuminate, almost glabrous. Pedicellate ovary 5 7 mm long, glabrous. Sepals 3-nerved. Dorsal sepal 2.7 3.6 mm long, 1.8 2.1 mm wide, oblong-ovate, obtuse. Lateral sepals 2.7 4.2 mm long, 2.0 2.3 mm wide, obliquely ovate, attenuate towards an obtuse apex. Petals 2.8 3.6 mm long, 0.8 1.1 mm wide, oblong-oblanceolate, obtuse, with glabrous margins. Lip up to 2.2 3.0 mm long, 2 3 mm wide, cochleate, suborbicular, subacute to obtuse at apex, with the midvein unbranched, but lateral veins branching. Gynostemium 0.9 1.1 mm long. Ecology Cranichis brevirostris grows at about 3000 3500 m a.s.l. Flowering throughout the year. Distribution Cranichis brevirostris is known from Colombia and Venezuela (Fig. 7). 6-EV

Figure 6. Comparison of the floral elements of (A) Cranichis silvicola, Renz 6139, RENZ and (B) C. ciliate, Silverstpone-Sopkin et al. 3817, CUVC). Drawn by A. Król and M. Kolanowska. 7-EV

long, 1.0 1.6 mm wide, narrowly elliptic-obovate, obtuse, 3-nerved. Lateral sepals 3 mm long, 1.3 mm wide, obliquely oblong-ovate, obtuse, 3-nerved. Petals 2.8 mm long, 0.4 0.5 mm wide, oblong-ligulate, obtuse, margins minutely glabrous or ciliate. Lip about 3 mm long, 2.4 mm wide, cochleate, lip from a cuneate base suborbicular to obovate, rounded to subacute at apex, with somewhat undulate margins, with the midvein branched and lateral veins branching. Gynostemium 1.5 mm long. Ecology Cranichis silvicola grows at about 1450 1750 m a.s.l. Flowering in November. Distribution Cranichis silvicola is known from Venezuela and Colombia (Fig. 7). Figure 7. Distribution of Cranichis badia (circle), C. brevirostris (square) and C. silvicola (triangle) in northern South America. Similar species Renz noted that Cranichis brevirostris is possibly only a variation of C. monophylla, however, it is easily distinguished from this species by the crenate leaf margins, the suborbicular lip lacking callosities (vs lip ovate-oblong with numerous fleshy projections on the inner surface), subdensely-flowered inflorescence (vs rachis laxly flowered) and floral bracts longer than ovary (vs floral bracts shorter than ovary) (Fig. 4). Additional specimens examined (paratypes) Colombia, Dept Cundinamarca, Plants with unopened flowers collected on 20 Jul 1941, which flowered in Bogota in mid-aug 1941, 3000 m a.s.l., O. Renz 4160 (RENZ!), Dept Cauca, 3000 3500 m a.s.l., Mar 1886, Lehmann 5885 (RENZ!). Venezuela. Est. Mérida, 2880 m a.s.l., 20 Nov 1949. Renz 6156 (RENZ!). Cranichis silvicola Renz ex Kolan. & Szlach. sp. nov. (Fig. 5) Similar to Cranichis ciliata (Kunth) Kunth from which it differs by the narrowly elliptic-obovate, obtuse dorsal sepal (vs dorsal sepal ovate, apiculate), lip cuneate at the base and floral bracts subequal to the ovary (vs lip sessile, floral bracts distinctly shorter than ovary). Type: Venezuela, Est. Mérida, 1450 m a.s.l., 17 Nov 1949, O. Renz 6139 (holotype: RENZ!). Description Plants up to 50 cm tall, erect. Leaves 2, gathered in a basal rosette, petiolate; petiole about 15 cm long, narrow, canaliculate; blade 6.8 9.1 cm long and 3.4 4.1 cm wide, ovate-lanceolate or elliptic, acute or shortly acuminate, cuneate at base. Scape up to 40 cm long; rachis up to 7.6 cm long, rather laxly many-flowered. Floral bracts 5 7 mm long, lanceolate, acuminate, glabrous. Pedicellate ovary 5 7 mm long, glabrous. Dorsal sepal 2.8 3.2 mm Similar species Cranichis silvicola resembles C. ciliata (Kunth) Kunth from which it differs mainly by the narrowly elliptic, obtuse dorsal sepal (vs dorsal sepal ovate, apiculate), lip cuneate at the base and floral bracts subequal to the ovary (vs floral bracts distinctly shorter than ovary). From other species of the C. ciliata-complex (e.g. C. atrata Schltr., C. polyblephara Schltr., C. sororia Schltr.) C. silvicola is distinguishable by the long-petiolate leaves (vs leaves short-petiolate) and lip that is cuneate at the base with somewhat undulate margins (vs lip sessile, margins non-undulate). In its floral characters the new species somewhat resembles C. antioquiensis Schltr., from which it is easily distinguished by the petiole twice or more as long as the leaf blade (vs petiole subequal to the blade). According to the original drawings by Renz, the margins of the petals are ciliate, although we found them glabrous on the duplicate specimen deposited at K (Lehmann 8505). Because other, both vegetative and floral characters are equal in both specimens of the same collection, we believe that the petals ciliation is not a stable feature of C. silvicola (Fig. 6). Additional specimens examined (paratypes) Colombia, Dept Cauca, 1750 m a.s.l., Lehmann 8505 (K!, RENZ! p.p.). Venezuela, Est. Mérida, 1500 m a.s.l., 18 Nov 1949. O. Renz 6141 (RENZ!). Acknowledgements The curators and staff of the cited herbaria are thanked for their kind hospitality and assistance during visits and for making specimens available on loan. We are grateful to Anna Król for preparing the illustrations. The research was supported by the Polish Ministry of Science and Higher Education (research grant no. 8124/ B/PO1/2011/40). References Acuña, J. 1939. Catalogo descriptivo de las Orquideas Cubanas. Bol. Ticn. Estac. Exp. Agron., Santiago de las Vegas 60: 48. 8-EV

Álvarez-Molina, A. and Cameron, K. M. 2009. Molecular phylogenetics of Prescottiinae s.l. and their close allies (Orchidaceae, Cranichideae) inferred from plastid and nuclear ribosomal DNA sequences. Am. J. Bot. 96: 1020 1040. Carnevali, G. and Ramírez-Morillo, I. M. 2003. Cranichis. In: Steyermark, B. et al. (eds), Flora of the Venezuelan Guayana. Vol. 7. Myrtaceae Plumbaginaceae. Miss. Bot. Gard. Press, p. 287. Chase, M. W. et al. 2003. DNA data and Orchidaceae systematics: a new phylogenetic classification. In: Dixon, K. W. et al. (eds), Orchid conservation. Nat. Hist. Publ., pp. 69 89. Christenson, E. A. 2004. Deux nouvelles Orchidees terrestres d Equateur. Richardiana 4: 134 138. Cribb, P. 2003. Cranichis. In: Pridgeon, A. M. et al. (eds), Genera Orchidacearum. Vol. 3: Orchidoideae, part 2, Vanilloideae. Oxford Univ. Press, p. 33. Dressler, R. L. 1981. The orchids: natural history and classification. Harvard Univ. Press. Dressler, R. L. 1993. Phylogeny and classification of the orchid family. Dioscorides Press. Garay, L. A. and Romero-González, G. 1999. Schedulae Orchidium II. Harv. Pap. Bot. 4: 475 488. McNeill, J. et al. (eds) 2006. International code of botanical nomenclature, Vienna Code. Reg. Veg. 146. Swartz, O. 1788. Nova genera species plantarum seu prodromus descriptionum vegetabilium maximam partem incognitorum quae sub itinere Indiam occidentalem annis 1783 1787. Acad. M. Swederi. Szlachetko, D. L. 1995. Systema orchidalium. Fragm. Florist. Geobot., suppl. 3: 1 152. Szlachetko, D. L. and Rutkowski, P. 2000. Gynostemia orchidalium 1. Acta Bot. Fenn. 169: 295 296, Fig. 362 364. 9-EV