VENEZUELA. Mérida: coleccionado en el bosque de la Laguna Negra, Páramo de Muchuchies, alt.: 3500 m, 25 Nov. 1943, Zoraida Luces de Febres 267 (holotype: VEN; isotype: MO (MO1086043! [image!]) fragm. ex VEN)

Briceño (2010) proposed the new combination of Calamagrostis meridensis for a taxon endemic to páramos of Venezuela. Briceño (2010) transferred the species from Agrostis to Calamagrostis based on the upper glume having 2–3 veins and anatomical characters such as all vascular bundles having a double sheath and without a notably angular shape and short cells over the veins can be solitary, in pairs or in short series. However, the species habit is noted as long rhizomatous to stoloniferous with geniculate culms, a habit unknown in Calamagrostis or Deschampsia, which are tufted or very-short rhizomatous and tussock-forming. This taxon is most likely a species of Podagrostis (Griseb.) Scribn. & Merr. based on the small spikelets (2.6–3.8 mm long), palea subequal to the lemma, awn often lacking, a rachilla extension that is usually absent or, when present, very short and glabrescent, a callus glabrous or rarely with scarce short hairs and short anthers 0.7–1 mm long. The number of veins of the upper glume has been considered as a distinguishing character to differentiate Podagrostis from Calamagrostis (e.g. Rúgolo de Agrasar 2012) but the recent discovery of Podagrostis colombiana Sylvester & Soreng (Sylvester et al. in press) from the Colombian Andes with well-developed lateral veins of the upper glume, large anthers and tussock-forming habit has shown these characters to be labile in Podagrostis.

ECUADOR. Pichincha: Crece en los pajonales de los montes Pichincha, Chimborazo y El Altar [occid. M. Pichincha, Tablahuasi], Aug. 1888, L. Sodiro 25/10 (lectotype, designated here: W (W1916-0037841 [image!]); isolectotype: W (W1916-0037840 [image!])).

Known only from the type collected in Ecuador, Calamagrostis sodiroana likely belongs to Deschampsia sensu Saarela et al. (2017), earlier to Calamagrostis sect. Deyeuxia subsect. Stylagrostis due to the rachilla internode being extended between the glumes and floret. This species bears affinities to Deschampsia parodiana and Deschampsia podophora due to its open panicle and straight awn that usually does not or only slightly surpasses the glumes and blades usually much shorter than the flowering culms (Sodiro 1930: 70; Zulma Rúgolo de Agrasar, pers. comm.).

This taxon was first noted as “Calamagrostis chaseae auct., non Luces” by Dorr et al. (2000 [2001]: 56) and subsequently called “Calamagrostis sp. A” in the Flora of Guaramacal (Venezuela): Monocotyledons (Dorr 2014) but its identity remains ambiguous and needs further study. The species habit is noted as stoloniferous, a habit not known from Calamagrostis or Deschampsia, which are tufted or very-short rhizomatous and tussock-forming. The mention of short spikelets (3–4.4 mm long), florets (appearing to) almost reach or equal the length of the glumes and 1-veined upper glumes in the brief description by Dorr (2014: 221) makes it possible that this taxon belongs to Podagrostis or Agrostis, although mention of a puberulent callus and presence of a fairly long awn (3.5–6 mm long) makes it less likely that this species is a Podagrostis.

ECUADOR. [without precise locality], 1859, Jameson s.n. [#59] (holotype: P (P00740364 [image!])).

Known only from the type collected in Ecuador, the identity of this taxon remains ambiguous. Calamagrostis spruceana bears similarities to C. macrophylla, C. secunda and C. macrostachya due to its open inflorescence, short hairs of the callus, rachilla hairs not reaching the apex of the palea and awn inserted slightly above the middle of the lemma (Sodiro 1930: 70; Zulma Rúgolo de Agrasar, pers. comm.).

ECUADOR. Crece en las pajonales del Pichincha entre 3650 y 4200 m, Sodiro s.n. (not located).

Known only from the type and the identity of this taxon remains ambiguous.

Calamagrostis pittieri, recorded for Costa Rica, Colombia and Venezuela (Morales Quirós 2003; Hokche et al. 2008; Giraldo-Cañas 2011, 2013; Giraldo-Cañas et al. 2016) is synonymised under Calamagrostis planifolia as no satisfactory characteristics were found to separate the two. Both have spikelets with florets bearing a pilose rachilla extension with hairs reaching from ¾ to passing the apex of the lemma, an awn inserted in the upper half of the lemma, 2 anthers that usually are short, ca. 1–1.4 mm long and leaf blades that are usually flat (sometimes drying convolute) with variable indumentum.

Calamagrostis pubescens was considered an endemic species to Colombia (Giraldo- Cañas 2013) and only known from the type specimen that was collected in southern Colombia from hills surrounding Pasto of departamento Narino. The B holotype was destroyed during World War II, with fragments only remaining at BAA and US. The BAA fragment consists of spikelets whose size and characteristics match C. planifolia (Zulma Rúgolo de Agrasar, pers. comm.), and the description of vegetative characteristics in the protologue also matches C. planifolia and so we consider C. pubescens a synonym of C. planifolia. Nevertheless, further study is needed for the C. planifolia complex in Colombia and there are certain crucial characters, such as number of stamens that need to be verified in these taxa. The C. pubescens BAA fragment lacked anthers (Zulma Rúgolo de Agrasar, pers. comm.) and Pilger (1898: 712) did not describe them.

Deyeuxia macrostachya is known only from the type specimen collected by Sodiro in páramo grasslands close to mount Pichincha and Chimborazo, Ecuador. We found no noticeable differences between this and Pilgers specimens. The large, lax panicle with semiverticillate branches, similar spikelet morphology including the lemma apex bifid with aristulate teeth and rachilla hairs only reaching the apex of the palea, support D. macrostachya being synonymised under C. macrophylla.

COLOMBIA. Magdalena: flanco occidental de la Sierra Nevada de Santa Marta, páramo, abundantisima, cubre gran parte del páramo, 3140 m alt., 29 Jan. 1959, R. Romero Castañeda 7141 (holotype: COL (COL000172001!); isotype: US (US01246667!)).

Figure 1. 

Calamagrostis crispifolius; A lower portion of plant, showing basal tuft of curled leaves B lígular area C upper portion of plant showing inflorescence and flag leaf blade D primary branch of the inflorescence E lower glume, dorsal view F upper glume, dorsal view G spikelet, with floret already fallen, lateral view H spikelet, with floret, lateral view I lemma, dorsal view J lemma, lateral view K stylar branch and stigma L palea ventral view, showing the ovary, stamens and lodicules M ovary and lodicules, ventral view N immature caryopsis, dorsal view showing embryo; drawn by Alice R. Tangerini from the isotype, R. Romero Castañeda 7141 (US).

Differs from all other species of Calamagrostis s.l. by a combination of strongly curled, readily deciduous leaf blades in mature plants that form a basal mat to 20 cm tall, open inflorescences with generally patent branches, spikelets (3.5–)4–5.5 mm long, with sessile florets and a rachilla prolongation (not including hairs) reaching from 2/3 to almost the apex of the lemma, with short hairs < 1 mm long and an awn inserted just above the middle of the lemma, 5–7.2 mm long, anthers 1.5–2.7 mm long.

Plants perennial, tufted, forming short dense tufts, mats to 40 cm wide, with short vertical or oblique rhizomes. Bases covered with fibrous old basal sheaths, with fallen curled leaf blades often forming large masses on the ground between tufts. Tillers intravaginal. Culms 48–64 cm tall, 1.5–2.2 mm wide, striate, erect, greatly exerted from the basal foliage, nodes and internodes terete, smooth but becoming scabrous below the nodes and towards the inflorescence, with dense scabrocities just below the inflorescence; (0–)1 node exposed at flowering; uppermost internodes 38–42 cm long. Sheaths striate, moderately keeled; flag leaf sheaths 20–25 cm long; upper culm sheaths glabrous and smooth with minute papillae present on the adaxial surface; basal leaf sheaths 4–12 cm long, shorter than the internodes, glabrous and lightly scabrous. Ligules not stipulate; upper culm ligules 2.5–10 mm long, acute with a rounded or bidentate apex, scarious to coriaceous, 2-veined but without notable lateral keels, apices denticulate, fimbriate or short ciliate, ligule abaxial surface lightly to densely scabrous with short scabers; ligules of innovations 2.2–10 mm long, strongly decurrent with the sheaths, truncate to acute, scarious to coriaceous when shorter, lightly to densely scabrous on the abaxial surface. Leaf blades 5–15(–30) cm long, 0.5–1.5 mm wide, cylindrical in outline, when rolled the blades form a basal mat to 20 cm tall and much shorter than the exerted culms, strongly curled in mature plants [or when dry?], appearing readily deciduous and snapping off when the plant reaches maturity leaving an abscission zone and the ligule exposed, sometimes recurved to straight in immature plants [or when moist?], conduplicate to convolute, rigid, glabrous, abaxially finely scabrous, adaxially densely scabrous, edges smooth or slightly scaberulous, apex pungent; flag leaf blade ca. 2.9 cm long, recurved, slightly narrower than the basal blades. Panicles (5.5–)9–20 cm long, 3–10 cm wide, open, rarely contracted in young specimens, exerted or rarely subincluded in the flag leaf, open and diffuse, oval, greenish-purple or golden-purple; main panicle axis terete, glabrous, lightly to moderately scabrous, spikelets found diffusely on the proximal half of the primary branches, lower internodes 2–4 cm long; panicle branches spreading to slightly ascending, rarely contracted; primary panicle branches 2–6 cm long, bearing 1–10 spikelets per branch, verticillate in clusters of 2–5, terete, glabrous, almost smooth to scabrous; pedicels (3–)6–22 mm long, usually much longer than the spikelets, glabrous, lightly to moderately scabrous. Spikelets 1-flowered, not strongly laterally compressed, disarticulating above the glumes; glumes, lemma and palea not noticeably asymmetrical. Glumes (3.5–)4–5.5 mm long, subequal, the lower glume ca. 0.3 mm shorter than the upper glume, narrowly lanceolate, membranous, purplish, lustrous, dorsal surface smooth or scaberulous distally, keels lightly scabrous distally or scabrous throughout, apices acute, finely denticulate, erose; lower glume 1-veined; upper glume 3-veined, lateral veins reaching from ¼ to past half the length of the glume, 1 or 2 cross veins between the keel and lateral vein infrequently present in ca. 10% of spikelets seen (requires 50× magnification). Floret sessile, almost reaching the apex of the glumes, sometimes passing the apex of the lower glume. Lowermost rachilla internode not prolonged between the glumes and the floret. Lemmas 3.7–4.6 mm long, 5-veined, veins not evident; the same consistency as the glumes, golden, glabrous, scaberulous throughout or lustrous and faintly to densely muriculate with the apex sometimes becoming scaberulous, apex shortly emarginate with lobes finely denticulate, awns 5–7.2 mm long, amply passing the glumes, twisted at the base, slightly curved, densely scabrous throughout, inserted just above the middle of the lemma, at 2.2–2.5 mm from the lemma base. Paleas 0.3–0.8 mm shorter than the lemma, of the same consistency and colour, keels smooth and notable, apex bidentate. Callus rounded, short, articulation oblique, with a basal tuft of short hairs 0.2–0.7(–0.8) mm long. Rachilla (2–)3–3.5 mm long, reaching from 2/3 to almost the apex of the lemma, with copious short hairs 0.5–1 mm long, the hairs reaching from 4/5 to almost the apex of the lemma and usually surpassing the palea. Lodicules 2, 0.4–0.6 mm long, membranous, 2-lobed, acute to slightly acuminate. Stamens 3, anthers 1.5–2.7 mm long. Ovary ca. 0.5 mm long, small, styles 2, stigmas plumose with secondary branching, short. Caryopsis ca. 1.8 mm long, ca. 0.7 mm wide, elliptic, rounded triangular in transection, hilum narrowly elliptic, ventral groove shallow and not conspicuous, pale brown, embryo ca. 0.3 mm long, apex with remains of styles and stigmas; endosperm firm.

Colombia, Venezuela. Known from páramos of the Sierra Nevada de Santa Marta, northern Colombia and the Sierra de Perija, Venezuela. For the Sierra Nevada de Santa Marta, specimens are known from páramo vegetation on both the western and eastern flanks of the massif. For the Sierra de Perija, specimens are known from both the northern and southern extents of the mountain range. It is found growing between 2700–3570 m in páramos with dry soils that are often subject to fires. The plant forms dense cushion-like mats or clumps and it is a dominant component of certain páramos. The type specimen label includes “Abundantisima, cubre gran parte del páramo”, i.e. highly abundant and covers a large part of the páramo. The specimen label of Barclay and Juajibuoy 6545 (US) also states “the dominant grass on these slopes, extending up to the top of ridge”, thus implying that it is a dominant component of the páramos of the Sierra Nevada de Santa Marta.

The degree of curling of leaf blades in the different specimens studied may relate to the level of maturity of the plant or also the local microclimate, with the specimen label of Barclay and Juajibuoy 6545 (US) stating “leaves very fine and rolled, when dry they curl”. The characteristic of readily deciduous leaf blades is also interesting, with the specimen label of Barclay and Juajibuoy 6545 (US) also mentioning that “large masses of these [the fallen curled blades] on the ground between clumps [of the plant]”.

COLOMBIA. Magdalena: Sierra Nevada de Santa Marta, Laguna Chubdula, 3480 m alt., 10°55'N; 73°53'W, 29 Dec. 1972, Kirkbridge & Forero 1775 (MO); Sierra Nevada de Santa Marta, alrededores de cabeceras del Río Sevilla, 3050–3300 m alt. [US specimen label states ‘The dominant grass on these slopes, extending up to the top of ridge. West and north facing slopes, on south side of river above campsite, sta.1,6., alt. 3320–3570 m’], 20 Jan. 1959, H.G. Barclay & P. Juajibioy 6545 (MO, US-3652630); Sierra de Perija, east of Manuare, Sabana Rubia, páramo, 3000–3100 m alt., 6 Nov. 1959, J. Cuatrecasas & R. Romero Castañeda 25021 (US [3 sheets]).

VENEZUELA. Zulia: Maracaibo Distr., Sierra de Perija, Serranía de Valledupar, Campamento Monte Viruela, on tepuí-like limestone massif 5×2.5 km in size, on the international boundary, 10°25.2167'N; 72°52.7'W, 3100 m alt., 25–28 Dec. 1974, S.S. Tillett 747-882 (MO); Perijá Distr., Sierra de Perija, Serranía de los Motilones, mesa below international boundary on main ridge, headwaters of Río Negro, Campamento Frontera II, 10°0.2167'N; 72°58.4167'W, 3000 m alt., 27 Nov.–5 Dec. 1974, S.S. Tillett & K.W. Hönig 746-618 (MO); Serranía de Valledupar, international boundary, headwaters of Río Guasare, 10°23.13'N; 72°52.0833'W, 2700–3300 m alt., 10–19 Dec. 1974, S.S. Tillett 747-1072 (MO).

Vulnerable (VU). Despite the species being known from seven collections spanning two Cordilleras, the páramos of Colombia are currently facing threats principally from mining (Pérez-Escobar et al. 2018) and an uncertain future. Our preliminary conservation status of VU is deemed adequate until further research is done.

The species epithet refers to the strongly curled leaf blades which make it distinct from all other páramo taxa of Calamagrostis s.l. with open panicles.

To our knowledge, there are no species of Calamagrostis s.l. with readily deciduous leaf blades that snap off when the plant reaches maturity leaving an abscission zone and the ligule exposed and covering the ground surrounding the plant tufts. This, coupled with the strongly curled nature of the leaf blades, makes this species unique. The character of curled leaf blades is very uncommon in the genus Calamagrostis, with the closest resembling species with this character being Calamagrostis crispa (Rúgolo & Villav.) Soreng, a species found in dry Andean grassland of Bolivia, Chile, Peru and Northeast Argentina (Rúgolo de Agrasar 2012: 193). The blades of mature plants of C. crispa are generally curved rather than strongly curled, as in C. crispifolius. Calamagrostis crispa also differs from C. crispifolius by the short, few-flowered, inflorescences that are included within the basal foliage, large spikelets with glumes 5–8 mm long and lemmas (4.4–)5–5.5 mm long, amongst other characters.

Calamagrostis crispifolius also shares certain similarities with C. effusa in terms of characters of the inflorescence i.e. the open panicles with verticillate panicle branches, the short glumes to 5.5 mm long, the awn being inserted in the upper half of the lemma and, most importantly, the long rachilla usually extending past the apex of the palea and bearing short hairs < 1 mm long. Calamagrostis crispifolius and C. effusa also share a peculiar character of cross veins between the lateral veins and keel of the upper glume, but these are only noticeable at 50× magnification in about 1 in 10 spikelets. A more exhaustive search for this character in other taxa within Calamagrostis s.l. should be done to check its exclusiveness to C. crispifolius and C. effusa. Calamagrostis crispifolius can be easily distinguished from C. effusa by the strongly curled leaf blades with pungent apices which form a basal mat to 20 cm high that is usually much shorter than the flowering culms, while C. effusa has stiffly erect blades with obtuse apices forming tussocks 40–60(–107) cm tall. Ligule characteristics also differ, with C. crispifolius having ligules 2.2–10 mm long with acute apices, while C. effusa has ligules 1–2 mm long with truncate apices. In a recent molecular analysis, Calamagrostis effusa was found to be sister to Chascolytrum Desv. (Saarela et al. 2017), possibly warranting its own generic placement. Further collecting of this new species, with a focus on molecular sampling, should be done to clarify its phylogenetic affinities.

Specimens of C. crispifolius from the Sierra Nevada de Santa Marta, Colombia, differ from Venezuelan specimens in a number of attributes and may represent a subspecies, although further collections and studies need to be made to confirm this. Colombian specimens have narrower leaf blades (0.5–0.75 mm wide), shorter ligules (2.2–4 mm long), usually longer anthers (to 2.7 mm long) and rachillas with short hairs that often reach the apex of the lemma. Venezuelan specimens have broader leaf blades (to 1.5 mm wide), longer ligules (4–10 mm long), shorter anthers (1.5–2 mm long) and rachillas with short hairs that usually do not reach the apex of the lemma.

COLOMBIA. Magdalena: Sierra Nevada de Santa Marta, valley descending south-western from Picos Reina and Ojeda, rocky and sandy páramos above Laguna Naboba and Laguna Reina, superpáramo, 4300–4500 m alt., 5 Oct. 1959, J. Cuatrecasas & R. Romero Castañeda 24607 (holotype: COL (COL000184738!); isotype: US (US01240776!)).

Figure 2. 

Deschampsia santamartensis. A Habit B Ligule C Panicle branch D Spikelet E spikelet, with lower glume pulled away to reveal the stipitate floret (extension of the lowermost rachilla internode) F floret G floret, with palea and lemma slightly separated to reveal the ovary in a late stage of development and surrounded by long filamentous trichomes H floret, with palea and lemma slightly separated to reveal the stamens and ovary at an earlier stage of development and surrounded by long filamentous trichomes; drawn by Marcela Morales from the holotype, J. Cuatrecasas & R. Romero Castañeda 24607 (COL).

Deschampsia santamartensis is similar to Deschampsia hackelii (Lillo) Saarela, but differs in having broad, rigid and erect, strongly conduplicate blades, 1.5–2.5 mm when folded (vs. filiform and curved leaf blades 0.5–1 mm wide when folded), ligules short and often truncate or obtuse with ligules of innovations 0.5–1 mm long and ligules of upper flowering culms 3–4 mm long (vs. ligules long and acuminate 3.5–7 mm long), ellipsoid spike-like panicles 3–5.5 long × 1.5–2.5 cm wide (vs. capituliform spherical panicles 1–3 cm long × 1–2 cm wide), lemma surfaces moderately to lightly scabrous between the veins (vs. smooth), lemma apices acute to muticous and entire (vs. truncate and irregularly dentate), rachilla extension often absent (vs. always present, 0.5–1 mm long) and inside of the floret often with hyaline shiny sinuous trichomes to 1 mm long emerging from the base of the ovary (vs. lacking trichomes).

Tufted perennial forming short dense tufts with vertical rhizomes and papyraceous, fibrous basal sheaths, leaves form a basal mat to 12 cm tall and much shorter than the exerted culms. Tillers intravaginal. Culms 9–20 cm tall, 1–1.5 mm wide, erect, exerted from the basal foliage, not obviously striate, densely papilliate; nodes and internodes terete, densely papilliate, not lustrous, nodes hidden in the sheaths with no nodes exposed at flowering; uppermost internodes 10–18 cm long, longer than the sheaths. Sheaths weakly striate, slightly keeled; flag leaf sheaths 7–10 cm long; upper culm sheaths lax, glabrous, smooth, densely papilliate; basal leaf sheaths 2–3 cm long, longer than the internodes, glabrous, smooth or slightly scabrous, densely papilliate, older basal sheaths papyraceous and fibrous with the fibres becoming slightly curly as the sheaths decay and leave just the fibres. Ligules not stipulate; upper culm ligules 3–4 mm long, regularly decurrent with the sheaths, broadly shouldered with an attenuate central point, hyaline, without notable lateral keels, apices entire or irregularly dentate, abaxial surface smooth and sparsely papilliate; ligules of innovations 0.5–1 mm long, strongly decurrent with the sheaths, broadly shouldered and sometimes with a lateral extension or lobe that extends ca. 1 mm long, the centre of the ligule truncate to obtuse and irregularly dentate, hyaline, without notable lateral keels, abaxial surface scabrous with distinct, mostly retrorse, spinules, with spinules tending to run down the throat and occur on the collar margin. Leaf blades 2.5–8 cm long, 1.5–2.5 mm wide when folded, gradually reduced up the culm, strongly conduplicate with margins frequently inrolled, weakly keeled, erect to slightly divergent, glabrous, abaxial blade surface smooth, strongly papilliate, adaxial blade surface scabrous mainly in the centre in lines along the veins, veins pronounced, margins moderately scaberulous, apex naviculate with a slightly pungent mucronate point; flag leaf blade 1.7–3 cm long, slightly narrower than lower culm blades. Panicles 3–5.5 cm long, 1.5–2.5 cm wide, dense, ellipsoid, golden-purple, densely spiculate, sometimes interrupted towards the base, spikelets present from near the base; main panicle axis terete, glabrous, smooth, lower internode ca. 1 cm long; panicle branches short, ascending; primary panicle branches 1–2 cm long, bearing 15–50 spikelets per branch, verticillate in clusters of 1–3, terete, glabrous, almost smooth to lightly scabrous, papilliate; pedicels 0.5–1.5 mm long, much shorter than the spikelets, glabrous, lightly scabrous, papilliate. Spikelets 1-flowered, strongly laterally compressed, disarticulating above the glumes (not seen); glumes, lemma and palea slightly asymmetrical. Glumes 4.6–5.5 mm long, subequal or rarely unequal, lower glumes 0.5–1.5 mm shorter than the upper glumes, membranous, purplish grading to a scarious golden-bronze margin, sub-lustrous, papilliate, very sparsely scabrous on the keels, edges smooth, apices acute to acuminate, entire to infrequently irregularly denticulate; lower glume 1-veined; upper glume 3 or sometimes faintly 5-veined, lateral veins reaching half the length of the glume. Florets stipitate, single, included in the glumes, subequalling the apex of the lower glume. Lowermost rachilla internode (0.25–)0.4–0.5 mm long, prolonged between the glumes and the floret, terete, slightly dilated at its apex, glabrous, smooth. Lemmas 4–4.5 mm long, of the same consistency as the glumes, golden-purple with a broad scarious margin and apex, glabrous, long scabrous to pectinate scabrous along the keel for most its length, lemma surface moderately to sparsely scabrous between the veins distally for up to ¾ the length of the lemma, apex slightly falcate, acute to muticous, entire, apex margins broadly scarious and somewhat incurved, 5-veined, veins generally not evident, sometimes apparent towards the base; awn absent. Paleas slightly shorter than the lemma, of the same consistency, golden, scarious, keels closely spaced with the keel flanges twice as broad as the gap between the keels, one keel more pronounced than the other, regularly scabrous and notable in the upper 2/3 of the length, scabrous between the keels, apex sometimes inconspicuously bidentate. Callus base rounded and slightly dorsally compressed above, glabrous. Rachilla absent or to 0.6(–1) mm long, glabrescent with a few short hairs at the apex. Lodicules 2, ca. 0.4 mm long, as broad as long, membranous, flabellate, with an acute lobe. Stamens 3, anthers 2–2.2 mm long. Ovary ca. 0.6 mm long, small, styles 2, stigmas plumose with secondary branching, short, often with scarce hyaline, shiny and sinuous trichomes to 1 mm long emerging from the base of the ovary. Caryopsis ca. 2 mm long, ca. 0.6 mm wide, obovate, laterally compressed in cross section, hilum 0.2 mm long, circular to obovate, ventral groove shallow and narrow and not conspicuous, pale honey brown, embryo ca. 0.4 mm long, apex with remains of style bases < 0.1 mm long, with remains of short plumose stigmas attached; endosperm liquid.

Known only from the type specimen that was collected from a southwest facing valley descending from Picos Reina and Ojeda in the centre of the Sierra Nevada de Santa Marta. The specimen was collected from high elevation rocky and sandy superpáramo vegetation above 4300 m.

Data Deficient (DD). Currently known only from a single specimen. Further expeditions are needed to the Sierra Nevada de Santa Marta to document its distribution.

The species epithet refers to the type locality of the Sierra Nevada de Santa Marta.

Deschampsia santamartensis clearly belongs to Calamagrostis subsect. Stylagrostis (=Deschampsia sensu Saarela et al. 2017) due to the presence of an extended rachilla internode between the glumes and the floret. Deschampsia santamartensis can be easily distinguished from all other members of Calamagrostis subsect. Stylagrostis with dense, spike-like panicles by its florets lacking awns, a glabrous callus and the absence of a rachilla extension (or rarely with a diminutive glabrescent rachilla extension). The only other member of Calamagrostis subsect. Stylagrostis with dense spike-like panicles, glabrescent callus, glabrescent short rachilla extensions and which occasionally lack awns is Deschampsia hackelii (Lillo) Saarela (= Calamagrostis hackelii Lillo), a species known from high-Andean regions of northwest Argentina and Chile (Rúgolo de Agrasar 2012: 201). Deschampsia hackelii differs by its capituliform panicles, 1–3 cm long × 1–2 cm wide (vs. ellipsoid spike-like panicles 3–5.5 long × 1.5–2.5 cm wide), filiform and curved leaf blades 0.5–1 mm wide when folded (vs. broad, rigid and erect, strongly conduplicate blades 1.5–2.5 mm when folded), long acuminate ligules 3.5–7 mm long (vs. ligules short and often truncate or obtuse with ligules of innovations 0.5–1 mm long and ligules of upper flowering culms 3–4 mm long), lemma surfaces smooth (vs. moderately to lightly scabrous between the veins), lemma apex truncate and irregularly dentate (vs. acute to muticous and entire), rachilla extension always present 0.5–1 mm long (vs. often absent) and inside of the floret lacking trichomes (vs. often with hyaline shiny sinuous trichomes to 1 mm long emerging from the base of the ovary).

Deschampsia aurea, a species originally described from Ecuadorian páramos but which also occurs in Peruvian Jalca vegetation, is another member of Calamagrostis subsect. Stylagrostis that has dense, spike-like panicles and spikelet lemmas that occasionally lose the thin weak dorsally-inserted awn (Zulma Rúgolo de Agrasar, pers. comm.). Deschampsia aurea has well-developed callus and rachilla hairs, with callus hairs surpassing more than half the length of the lemma and rachilla hairs reaching or surpassing the lemma apex, as well as other characters to help differentiate it from D. santamartensis such as the floret being noticeably shorter (2.9–3 mm long) than the large glumes (5.5–7.5 mm long), amongst other characters. Calamagrostis chrysostachya (E. Desv.) Kuntze, a species known from high-Andean regions of northwest Argentina and Chile, also shares these attributes and has also been included as a member of Calamagrostis subsect. Stylagrostis (Rúgolo de Agrasar 2012, pers. comm.). Both these species, however, have a pilose callus and rachilla, with C. chrysostachya having poorly developed callus and rachilla hairs to 1 mm long, but thin conduplicate or convolute leaf blades 0.8–1.2 mm wide when unfolded and a ligular stipule 0.5–1.5 mm long. The short, strongly conduplicate leaf blades with navicular apices are also reminiscent of Poa trachyphylla Hack. which also inhabits high-elevation superpáramo of Colombia (Sylvester et al. in press).

PERU. Junín: Berge Westlich von Huacapistana [Prov. Tarma, in montibus prope Huacapistana ad occid. in stepposis], 3500 m alt., 18 Jan. 1903, A. Weberbauer 2231 (lectotype, designated by Vega and Rúgolo de Agrasar (2013: 28): BAA (BAA00000767 [image!]) fragm. ex B; isolectotype: US (US00149282!)).

Tufted perennial with vertical rhizomes forming short solitary tufts to medium-sized tussocks, with leaf blades mostly basal with inflorescences usually greatly exerted from basal foliage or both basal and cauline with some cauline blades often surpassing the inflorescence. Tillers intravaginal. Culms 20–75(–110) cm tall, to 3 mm wide, erect, striate, nodes and internodes terete, smooth and lustrous; nodes hidden in the sheaths with no nodes exposed at flowering; uppermost internodes 20–32.5 cm long, as long or longer than the sheath. Sheaths striate; flag leaf sheaths 22–38 cm long; upper culm sheaths lax, glabrous and smooth; basal leaf sheaths 4.5–20 cm long, longer than the internodes, glabrous and smooth. Ligules not stipulate; upper culm ligules 7.5–22 mm long, strongly decurrent with the sheaths, long acuminate, membranous to slightly coriaceous, without notable lateral keels, apices entire, erose or narrowly bifid, sometimes fimbriate, abaxial surface smooth; ligules of innovations 4–15(–20) mm long, slightly to strongly and broadly decurrent with the sheaths, long acuminate, membranous to slightly coriaceous, lateral keels sometimes notable, apices entire or a narrow bifid point, sometimes slightly erose, abaxial surface smooth or sometimes slightly scabrous at the apex. Leaf blades (2.5–)5–22 cm long, 0.6–7 mm wide when opened out, flat, conduplicate or involute/convolute and filiform and cylindrical to subelliptical in outline, sometimes opening out to become flat at their apices, straight and erect to slightly curved, glabrous, isomorphic or more or less dimorphic, when dimorphic those of the innovations filiform and cylindrical to subelliptic in outline while those of the upper flowering culm are usually wider and flat, conduplicate or convolute towards the apices, abaxially smooth, adaxially smooth or lightly scaberulous along the veins, sometimes becoming densely scabrous towards the apex, edges smooth or slightly scaberulous, veins usually pronounced, numerous and tightly packed, apex obtuse to slightly pungent; flag leaf blades 2.9–6 cm long, 2–7 mm wide when opened out. Panicles 10–25 cm long, 3–8 cm wide, open to slightly condensed, oval, greenish-purple with spikelets tending to be laxly glomerate on the distal half of the inflorescence branches with the proximal half usually lacking spikelets, largely exerted to moderately included in the uppermost sheath and/or blade; main panicle axis terete to slightly compressed, usually with a narrow groove running down both sides, glabrous, smooth to lightly scaberulous, internodes tending to be long, lower internode 3–11.5 cm long; panicle branches 1.5–8 cm long, bearing 10 to over 50 spikelets per branch, flexuous, spreading, pendulous or divergent at a 45° angle to slightly ascending, verticillate in clusters of 2 or 3, terete or slightly grooved, glabrous, almost smooth to scabrous; pedicels 0.5–2.5 mm long, usually shorter than the spikelets, glabrous, lightly to densely scabrous. Spikelets 1-flowered, sometimes with a rudimentary floret at the apex of the rachilla that appears like a slightly broader section of the rachilla covered in sparse diminute hairs, not strongly laterally compressed, disarticulating above the glumes with the florets disarticulating from the apex of the extended rachilla internode, this remaining attached to the glumes. Glumes 3.5–5.5 mm long, subequal, the lower glume 0.2–0.6 mm shorter than the upper glume, lanceolate, membranous, purplish-green, lustrous, smooth or sometimes lightly scabrous throughout the keel of the upper glume; lower glumes 1-veined, apex acuminate or bidentate, less frequently finely denticulate or erose; upper glumes 3-veined, lateral veins either short < ½ length of glume or reaching from ½ to 2/3 the length of the glume, apex usually acuminate, sometimes finely denticulate. Floret stipitate, much shorter than the glumes, never passing the apex of the lower glume. Lowermost rachilla internode 0.4–0.7 mm long, prolonged between the glumes and the floret, often slightly geniculate at its apex and bent in a ca. 30°–45° angle, slightly dilated at its apex, usually glabrous, less often with a few long hairs ca. 0.7 mm long emerging from it, smooth. Lemmas 2.4–3.5 mm long, of the same consistency as the glumes, light green with purple tinges towards apex, becoming golden at maturity, glabrous, smooth with the keel apex rarely scaberulous, apex truncate and denticulate, usually with 4 clearly distinguished teeth, 0.3–0.5 mm long and erose between the teeth, 5-veined, veins not evident; awns 1.5–4 mm long, sometimes absent, inserted in the middle or lower third of the lemma, usually as long as the lemma or passing the glume apex by as much as 1.5 mm. Paleas 0.5–0.8 mm shorter than the lemma, of the same consistency and colour, keels sparsely scabrous and notable, apex bidentate or 4-dentate. Callus rounded, short, with a basal tuft of hairs 0.5–2.2 mm long, reaching from 1/3 the length of the lemma to almost the lemma apex. Rachilla 1.5–2.5 mm long, reaching from 2/3 to 4/5 the length of the lemma, with copious short to medium-sized hairs 0.5–1.4 mm long, the hairs reaching from 4/5 to sometimes passing the apex of the lemma, apex of rachilla sometimes clavate. Lodicules 2, ca. 0.5 mm long, membranous, acute. Stamens 3, anthers (0.7–)1.2–1.9 mm long. Ovary ca. 0.5 mm long, small, styles 2, stigmas plumose, short. Caryopses 1–1.4 mm long, dorsally slightly gibbose, surcus not noticeable, embryo short, hilum basal, oval; endosperm dry.

Colombia, Ecuador, Peru, Venezuela. The ecology of this species is distinct compared to many Calamagrostis s.l. species as it is usually found in very damp, swamp-like conditions by the side of high-elevation lakes or watercourses in Andean páramo or jalca vegetation, less often in humid open páramo.

COLOMBIA. Boyacá: Below Las Playas de Ritacuba in the Cocuy mountains above Guican, on the banks of a fast-flowing stream coming off the snow fields and passing through thinly vegetated moraine covered country, 4100 m alt., 24 Jun. 1984, J.R.I. Wood 4457 (K). Cauca: Purace National Park, Laguna de San Rafael, in open boggy páramo in the lake basin, particularly in banks by ditches, 3300 m alt., 6 Apr. 1985, J.R.I. Wood 4803 (K); Volcan Purace, above Pilimbala, frequent in bog pools in high páramo, 3700–4000 m alt., 5 Apr. 1985, J.R.I. Wood 4787 (K).

ECUADOR. Pichincha: road Olmedo-Laguna San Marcos, W of the pass, 0°5'N; 78°1–2'W, 3600 m alt., 10 Jul. 1980, B. Øllgaard et al. 34406 (K); Along road to Refugio, Volcan Cayembe, páramo and swamp, 00°04'S; 77°54'W, 4300 m alt., 2 Mar. 1988, S. Renvoize 70510 (K). Napo: Eastern Cordillera, Llanganati Mountains, by Lake Aucacocha, on stream sides in bog, forming large tussocks up to 20 cm across, 3750 m alt., 16 Aug. 1969, P.J. Edwards 127 (K); Eastern Cordillera, Llanganati Mountains, by Lake Aucacocha, growing in the wettest area of the bog, in clusters of tufts, sheath bases submerged in peat, 3700 m alt., Aug. 1969, P.J. Edwards 62 (K).

Deschampsia podophora has been traditionally treated as belonging to Calamagrostis subsect. Stylagrostis due to the presence of an extended rachilla internode between the glumes and the floret. Deschampsia podophora is closely related to D. parodiana (=Calamagrostis ligulata) and has been placed as a synonym of this in previous works (Escalona 1988a, 1988b; Tovar 1993; Luteyn 1999; Bono 2010; Briceño 2010). The principal differentiating characters that separate Deschampsia parodiana from D. podophora are the smaller anthers, 0.4–0.5 mm long (vs. (0.7–)1.2–1.5(–1.9) mm long in D. podophora) and the shorter rachilla, 1–1.2 mm long, that is sparsely pilose with hairs not usually reaching the apex of the palea (vs. (1.2–)1.4–2.5 mm long, with copious hairs that usually surpass the lemma in D. podophora). Characters of shape and density of the inflorescence mentioned in Laegaard (1998: 27) were found to be not good for differentiating the two species. While D. parodiana only has a lax open inflorescence with long pendant branches and spikelets glomerate on the distal part of the branches, D. podophora exhibits both denser semi-spikelike inflorescences with inflorescence branches having spikelets from the base as well as open inflorescences with pendulous branches and spikelets glomerate on the distal part of the branches like that of D. parodiana.

Some specimens from Colombia were found to be generally larger than those from Ecuador, in terms of the number and length of the culms with mainly cauline leaf blades that were longer and wider. These specimens had a habit appearance similar to var. mutica being larger tussock-forming plants with multiple culms and inflorescences held within sheaths.

COLOMBIA. Cundinamarca: Mun. Santa Rosa-Usme, Sumapaz páramo, by Laguna Larga, in swamp at the edge of the lake in open páramo country, 3700 m alt., 19 Aug. 1985, J.R.I. Wood 5033 (holotype: FMB (FMB11918!); isotypes: COL (COL000434793!), K [2 sheets]!).

Figure 3. 

Deschampsia podophora var. mutica. A whole plant B anther C spikelet at maturity with anthers D lemma, abaxial view E palea, abaxial view F upper glume, abaxial view G lower glume, abaxial view H floret I spikelet and pedicel, lateral view J ligule; drawn by Juliet Beentje from the isotype, J.R.I. Wood 5033 (K).

Deschampsia podophora var. mutica differs from D. podophora by the lemmas being muticous and lacking awns (vs. lemmas usually with a well-developed dorsal awn inserted in the lower or middle third of the spikelet, measuring 1.5–3.5(–4) mm long and usually not surpassing the glumes), tussock-forming habit with multiple culms and leaves mainly cauline, without a basal mat clearly shorter than the flowering culms (vs. plants forming short isolated tufts with solitary culms and the leaves forming a short basal mat clearly shorter than the largely exerted flowering culms), inflorescences often sub-included in the sheaths and blades (vs. flowering culms largely exerted from basal mats); leaf blades 11–22 cm long, 0.35–0.6 mm wide when rolled, often dimorphic, those of the innovations filiform and cylindrical to subelliptic in outline, while upper flowering culm blades 6.5–25 cm long, 2–7 mm wide when opened out, usually wider and flat, conduplicate or convolute towards the pungent apices (vs. leaf blades 5–8 cm long, 2–3 mm wide, not clearly dimorphic, with all blades flat or conduplicate, apices obtuse), ligules 4–11 mm long (vs. ligules generally longer, 10–22 mm long), anthers (1.4–)1.8–1.9 mm long (vs. anthers (0.7–)1.2–1.5 mm long), upper glume lateral veins reaching from ½ to 2/3 the length of the glume (vs. upper glume lateral veins short, < ½ length of glume).

Tufted perennial with vertical rhizomes forming medium-sized tussocks with leaf blades both basal and cauline and some cauline blades often surpassing the inflorescence. Culms 26–110 cm tall, to 3 mm wide, erect, striate, nodes and internodes terete, smooth and lustrous; nodes hidden in the sheaths with no nodes exposed at flowering; uppermost internodes 23–32.5 cm long, not, or not pronouncedly, longer than the sheath; Sheaths striate; flag leaf sheaths 22–38 cm long; upper culm sheaths lax, glabrous and smooth; basal leaf sheaths 4.5–20 cm long, glabrous and smooth, longer than the internodes. Ligules not stipulate; upper culm ligules 7.5–11 mm long, strongly decurrent with the sheaths, long acuminate, membranous to slightly coriaceous, without notable lateral keels, apices erose or narrowly bifid, sometimes fimbriate, abaxial surface smooth; ligules of innovations 4–9 mm long, slightly to strongly decurrent with the sheaths, long acuminate, membranous to slightly coriaceous, lateral keels sometimes notable, apices a narrow bifid point, sometimes slightly erose, abaxial surface smooth or sometimes slightly scabrous at the apex. Leaf blades sometimes dimorphic, those of the innovations filiform and cylindrical to subelliptic in outline while those of the upper flowering culm are usually wider and flat, conduplicate, or convolute towards the apices; leaf blades of innovations 11–22 cm long, 0.35–1 mm wide when rolled or folded, narrow and conduplicate or filiform involute or convolute and cylindrical to subelliptical in outline, rarely completely flat, sometimes opening out to become flat at their apices, straight and erect, glabrous, abaxially smooth, adaxially lightly scaberulous along the veins or rarely smooth, edges smooth or slightly scaberulous, apex obtuse to slightly pungent; leaf blades of lower flowering culm to 1.2 mm wide when rolled or folded, similar to those of the innovations or slightly wider; leaf blades of upper flowering culm 6.5–25 cm long, 2–7 mm wide when opened out, flat, conduplicate or convolute towards the apices, glabrous, abaxially smooth to finely scaberulous, sometimes becoming densely scabrous at the apex, adaxially smooth or scaberulous towards the margins, veins pronounced, numerous and tightly packed, edges smooth or slightly scaberulous, apex acute to pungent; flag leaf blade ca. 2.9 cm long, recurved, slightly narrower than the basal blade. Panicles 10–25 cm long, 5–8 cm wide, open and diffuse with main axis having long internodes, oval, usually slightly to moderately included in the uppermost sheath and/or blade, greenish-purple, spikelets tending to be laxly glomerate on the distal half of the inflorescence branches with the proximal half usually lacking spikelets; main panicle axis terete to slightly compressed, usually with a narrow groove running down both sides, glabrous, smooth to lightly scaberulous, internodes tending to be very long, lower internode 5.5–11.5 cm long; panicle branches flexuous, spreading, pendulous or divergent at a 45° angle to slightly ascending; primary panicle branches 1.5–8 cm long, bearing 10 to over 50 spikelets per branch, terete and slightly grooved, verticillate in clusters of 2 or 3, glabrous, almost smooth to lightly scabrous; pedicels 0.5–2.5 mm long, usually shorter than the spikelets, glabrous, lightly scabrous. Glumes 4.5–4.9 mm long, subequal, the lower glume 0.3–0.6 mm shorter than the upper glume, membranous, purplish-green, lustrous, smooth or sometimes lightly scabrous throughout the keel of the upper glume; lower glume 1-veined, apex usually bidentate, less frequently finely denticulate or erose; upper glume 3-veined, lateral veins reaching from ½ to 2/3 the length of the glume, apex usually acuminate, sometimes finely denticulate. Floret stipitate, much shorter than the glumes, never passing the apex of the lower glume. Lemmas 2.8–3.4 mm long, of the same consistency as the glumes, light green with purple tinges towards apex, becoming golden at maturity, glabrous, smooth with the keel apex rarely scaberulous, apex truncate and denticulate, usually with 5 clearly distinguished teeth, 0.3 mm long, and erose between the teeth, 5-veined, veins not evident; muticous and lacking an awn. Rachilla 1.5–2.5 mm long, reaching from 2/3 to 4/5 the length of the lemma, with copious short to medium-sized hairs 0.5–1.4 mm long, the hairs reaching from 4/5 to almost the apex of the lemma and usually surpassing the palea, apex of rachilla often clavate. Stamens 3, anthers (1.4–)1.8–1.9 mm long.

Endemic to Colombia. Known from páramos of the Cordillera Oriental and Cordillera Central of the Colombian Andes. For the Cordillera Oriental, the species is known from Departamento Cundinamarca municipalities Usme and Santa Rosa and Páramo Pisba of Departamento Boyacá. For the Cordillera Central, the species is known from Páramo del Quindio and Páramos de la Laguna del Mosquito of Departamento Caldas. Found in humid, swampy areas, often by rivers or lakes and less often in more mesic habitats, such as road verges (presumed damp). The type specimens were collected from swampy areas bordering the Laguna Larga of the Sumapaz páramo in Cundinamarca. The holotype shows signs of grazing, with blades and culms abruptly cut.

COLOMBIA. Boyacá: Mun. Socota: Páramo Pisba, Peña Negra, lagoon “Choro Negro”, páramo with Calamagrostis effusa, Espeletia sp., Chusquea sp., Diplostephium sp. etc., swampy, 3500 m alt., 11 Feb. 1999, D. Stančik & S. Medina 2351 (COL-000184768; FMB-051200); Páramo Pisba, Alto de Calarca, humid grassy páramo with Calamagrostis effusa, Espeletia sp., Chusquea sp. et bunch Grass, 3600 m alt., 11 Feb. 1999, D. Stančik & S. Medina 2331 (FMB-046617). Caldas: Páramo del Quindio, swale in páramo valley, 3700–4200 m alt., 15–20 Aug. 1922, F.W. Pennell & T.E. Hazen 9949 (K); Cordillera Central, cabeceras del río Otúm, bajando del Nevado de Santa Isabel, páramos de la Laguna del Mosquito, 3820 m alt., 26 Nov. 1946, J. Cuatrecasas 23233 (K). Cundinamarca: Mun. Usme: Laguna Chizaca, hierba creciendo al lado de la carretera que conduce a la laguna, 26 Jul. 1986, A. Betancur & M. Palacio 44 (HUA-47083); Páramo de Chisaca, growing in marsh near lagoon, 3750 m alt., 30 Sep. 1966, T.R. Soderstrom 1273 (K).

Vulnerable (VU). Despite the species being known from several collections from the Central and Eastern Cordilleras of the Colombian Andes, the páramos of Colombia are currently facing threats of habitat degradation and loss, principally from mining (Pérez-Escobar et al. 2018) and so a preliminary conservation status of VU is given.

The varietal epithet refers to the absence of an awn inserted in the dorsal surface of the lemma.

Certain specimens were encountered which exhibited a mix of the characters mentioned in the diagnosis for separating D. podophora from D. podophora var. mutica. For example, specimen Cuatrecasas 23233 from the Cordillera Central of Colombia had isomorphic flat leaf blades which formed a basal mat much shorter than the exerted culms but also had awnless spikelets. A few other specimens exhibited the larger habit, i.e. multiple longer culms with longer and wider leaf blades forming tussocks and inflorescences included within sheaths, but with awned spikelets.

Specimens of Deschampsia podophora var. mutica were commonly misidentified as Poa L. in herbaria, most likely due to the lemmas lacking awns. There are very few South American Calamagrostis s.l. that consistently lack awns (see ‘notes’ of Deschampsia santamartensis sp. nov. above) and, of the species not belonging to subsect. Stylagrostis (=Deschampsia), the closest resembling species is Calamagrostis ecuadoriensis Laegaard (1998), which also has short florets with a pilose rachilla extension and which lack awns. However, C. ecuadoriensis can be differentiated from Deschampsia podophora var. mutica by, amongst other things, a) habit, being small tussocks, 20–30 cm high, with leaves mostly basal; b) culms, panicle branches and pedicels densely hispid; c) panicles narrow, ca.1 cm wide; d) anthers 0.8–1 mm long; e) florets only shortly stipitate, with the stipe < 0.15 mm long at the base of the glumes.

ECUADOR. Napo [Sucumbíos]: Páramo de Mirador above Cocha Seca, lower páramo zone, burned, 00°34'N; 77°39'W, 3700–3900 m alt., 23 May 1985, S. Lægaard 54413 (holotype: AAU!; isotypes: K (K000308461!), MO (MO05100301 [image!]), QCA (QCA78857 [image!]), QCNE, US (US00588939!)).

Figure 4. 

Calamagrostis cf. carchiensis Lægaard general habit (scale bar 5 cm) and spikelet. Scale bar: 1 mm; S.P. Sylvester 3049 (FMB).

Previously considered endemic to Ecuador with a global conservation status of VU B1ab(iii) - Vulnerable (León Yánez et al. 2011). The voucher specimen collected matches the species description in every aspect apart from its having two anthers as opposed to one. The number of anthers is taxonomically informative in the genus Calamagrostis and it may be that the Sylvester 3049 specimen should be considered as a distinct taxon, although further research including molecular analysis is needed to clarify this. Laegaard (1998) noted that Calamagrostis carchiensis bears affinity to Calamagrostis bogotensis (Pilg.) Pilg., especially in terms of florets with a single anther.

COLOMBIA. Boyacá: Mun. Duitama, páramo de Agueros, via que conduce a Vereda de Avendanos, 05°54.527'N; 73°03.761'W, 3445 m alt., 28 Oct. 2017, S. P. Sylvester, W. Bravo & J. Aguilar 3049 (COL, FMB, K, US).

ECUADOR. Napo: [road Quito-Baeza at the telecomunication antenna, N of the Guamani paramo, in the oriental Andes, 0°10.2'S; 78°23.4'W], 4260 m alt., 3 Mar. 1985, [grass forming loose tufts in cushion plants of Distichia muscoides], F.D. Escalona & Gallegos 390 (holotype: ISC; isotypes: K!, MO (MO115961), QCA, US!, VEN).

Previously considered endemic to Ecuador (Luteyn 1999).

COLOMBIA. Nariño: Mun. Pasto, Volcan Galeras, frequent growing in cushion plants and similar damp protected places at high altitudes in base high páramo, 1°13.6417'N; 77°21.8718'W, 4000 m alt., 29 Nov. 1983, J.R.I Wood 4064 (FMB, K).

Figure 5. 

A general habit and a spikelet of Calamagrostis guamanensis Escalona, J.R.I. Wood 4064 (FMB), Calamagrostis heterophylla (Wedd.) Pilg., S.P. Sylvester 3158 (US), Calamagrostis pisinna Swallen, S.P. Sylvester 3107 (US), and Calamagrostis rigida (Kunth) Trin. ex Steud., S.P. Sylvester 3125 (US). Habit scale bar 5 cm; spikelet scale bar 1 mm.

Previously known from high Andean regions of Venezuela (Hokche et al. 2008; Bono 2010; Briceño 2010), Peru (Tovar 1993), Bolivia (Villavicencio 1995, 1998), northern Chile and northwest Argentina (Rúgolo de Agrasar 2012). The specimens collected exhibit characters of both Calamagrostis heterophylla and Calamagrostis brevipaleata Swallen, an Ecuadorian endemic, with both species having heteromorphic leaf blades, the cauline glabrous and those of the innovations pilose. The species bear more affinity to C. heterophylla as the lemmas measure less than 4.2 mm (> 5 mm long in C. brevipaleata) and the leaf blades measure < 10 cm long (10–25 cm long in C. brevipaleata). However, the specimens do have characteristics of C. brevipaleata in terms of the lemma surfaces, which are smooth proximally and scabrous distally and no great differentiation in width between the cauline and tiller leaf blades.

COLOMBIA. Boyacá: Mun. Chiscas, páramo El Penon, borde de bosque de Polylepis creciendo sobre roca, 6°36.0714'N; 72°26.229'W, 3917 m alt., 5 Mar. 2018, S.P. Sylvester, R.J. Soreng, W. Bravo & L.E. Cuta 3158 (COL, FMB, K, US).

Venezuela. Mérida: rocky ridges, higher paramos, near El Gavilon, 4200 m alt., 25 Jan. 1929, H. Pittier 13277-1/2 (lectotype, designated here: US (US00149283! [A-two flowering culms are the type, B-on left side of sheet is unknown])).

Previously considered endemic to Venezuela (Escalona 1988a) but specimens have been found in the Sierra Nevada del Cocuy of the Cordillera Oriental of Colombia. Specimen Sylvester 3107 differed slightly from the species described from Venezuela in that the leaf blades were densely pilose and often found to be conduplicate (Fig. 5E, F). As only one flowering specimen was encountered of this morphotype and specimens being found growing out of fairly inaccessible crag ledges, more collections need to be made to ascertain whether this may be a distinct species. All specimens from the Sierra Nevada del Cocuy have a rachilla extension with hairs that reach or surpass the apex of the floret, while Venezuelan specimens have rachilla hairs which usually do not surpass the apex of the palea (Escalona 1988a). Saarela et al. (2017) found one of the specimens of C. pisinna that we cite here (Cleef 8653) to have an unusual placement in plastid analyses (no nuclear ribosomal data was obtained) as a basally diverging lineage in a moderately to strongly supported clade that also contained Lagurus ovatus L., Aveninae s.str., and Koeleriinae excluding L. ovatus. However, there is a phylogenetic discrepancy between matK and PsbK sequences reported by Saarela et al. (2017) for C. pisinna that needs to be evaluated before any taxonomic conclusions can be drawn.

There is a sterile tuft on the left side of the Pittier type sheet that is a different species and quite possibly a different genus, having acicular involute blades and longer acute ligules that are decurrent (p.p. b). Thus, we lectotypify to the two flowering tufts on the right side of the type sheet (p.p. a), which are identical and fit the description in the type protologue.

COLOMBIA. Boyacá: Mun. Chiscas, páramo de Chacaritas, limit between páramo and superpáramo, growing out of rock ledge, 6°37.6794'N; 72°23.6616'W, 4072 m alt., 4 Mar. 2018, S.P. Sylvester, R.J. Soreng, W. Bravo & L.E. Cuta 3107 (US); Sierra Nevada del Cocuy, Páramo Cóncavo, Cueva de los Hombres, 3 km N del Morro Púlpito del Diablo, 4350 m alt., 28 Feb. 1973, A. Cleef 8610 (US-01234789); 4350 m alt., A. Cleef 8653 (US-01234736).

Previously considered to have its northernmost distribution in Ecuador (Rúgolo de Agrasar 2012), although Luteyn (1999) mentions its presence in Costa Rica, this is the first record of C. rigida for Colombia. It has been erroneously determined as Calamagrostis recta (Kunth) Trin. ex Steud. The latter differs from C. rigida by the rachilla hairs reaching up to ¾ the length of the lemma (as opposed to usually reaching to slightly surpassing the lemma apex in C. rigida), the awns being generally longer (6.2–7.5 mm as opposed to 4–6 mm in C. rigida) and only slightly surpassing the glumes, the lemma apex being slightly bidentate (as opposed to bifid in C. rigida) and the ligule being generally truncate and shorter, 1–5(–6.6) mm long (as opposed to long acuminate, (3–)8–12 mm long in C. rigida), amongst other things.

There has been discrepancy regarding the typification of Agrostis antoniana, with previous research (e.g. Rúgolo de Agrasar 2012) citing the specimen P.G. Lorentz and G.H.E.W. Hieronymus 72 housed at CORD herbarium from Argentina as holotype, while the protologue mentions this (although not explicitly giving collector and number) as well as three other collections: Spruce pl. ecuad. 5927 presumably from Ecuador, G. Mandon 1308 from Bolivia [Peru] and W. Lechler 1800 from Peru. A further specimen, P.G. Lorentz and G.H.E.W. Hieronymus 67, makes a total of at least five syntypes for this name, although the three GOET Lorentz and Hieronymus syntypes were annotated by Grisebach with “67 + 72”. Hitchcock (1927) gave a partial lectotypification by dictating the type as Lechler 1800, but did not indicate the herbarium. The Lechler 1800 specimen was distributed with the following label annotated by Hohenacker: “W. Lechler Pl. Peruvian. Ed. R. F. Hohenacker 1800 Calamagrostis Antoniana Steud. Ipse [“he himself said it”]. In graminosis pr. San Antonio rara Jun. m.”. As no annotations by Grisebach were found on any of the Lechler 1800, Mandon 1308 or Lorentz and Hieronymus 67 and 72 CORD specimens (Spruce pl. ecuad. 5927 specimens not found), with only Lechler 1800 specimens at P being verified by Steudel, we lectotypify to the best of the three GOET Lorentz and Hieronymus 67 + 72 syntypes annotated by Grisebach and consider syntypes Lorentz and Hieronymus 67 and Lorentz and Hieronymus 72 to be isolectotypes.

COLOMBIA. Boyacá: Mun. Chiscas, páramo de Chacaritas, asociado a rocas de 4 m de altura, 6°37.3362'N; 72°23.424'W, 4192 m alt., 4 Mar. 2018, S.P. Sylvester, R.J. Soreng, W. Bravo & L.E. Cuta 3119 (COL, FMB, US, UPTC); Mun. Chiscas, páramo de Chacaritas, arribando a la morrena, 6°37.0674'N; 72°23.3394'W, 4354 m alt., 4 Mar. 2018, S.P. Sylvester, R.J. Soreng, W. Bravo & L.E. Cuta 3125 (COL, FMB, K, SI, UPTC, US); Mun. Chiscas, páramo de Chacaritas, arribando a la morrena, 6°37.0674'N; 72°23.3394'W, 4354 m alt., 4 Mar. 2018, S.P. Sylvester, R.J. Soreng, W. Bravo & L.E. Cuta 3126 (US). Dep. Santander: Páramo de la Angostura, Vereda El Mortino, Ubicada en borde de quebrada, 6°57.5'N; 72°43.5'W, 3605 m alt., 17 Nov. 2007, M.C. Gomez 1 (US).