A revision of Poa subsection Aphanelytrum (Poaceae, Pooideae, Poaeae, Poinae)... 107
A revision of Poa subsection Aphanelytrum (Poaceae, 
Pooideae, Poaeae, Poinae); and a new species, 
Poa auriculata
Paul M. Peterson1, Robert J. Soreng1
1 Department of Botany MRC-166, National Museum of Natural History, Smithsonian Institution, Washington, 
DC 20013-7012 USA
Corresponding author: Paul M. Peterson (peterson@si.edu)
Academic editor: C. Morden    |   Received 20 February 2016    |   Accepted 25 May 2016    |   Published 6 June 2016
Citation: Peterson PM, Soreng RJ (2016) A revision of Poa subsection Aphanelytrum (Poaceae, Pooideae, Poaeae, Poinae); 
and a new species, Poa auriculata. PhytoKeys 63: 107–125. doi: 10.3897/phytokeys.63.8198
Abstract
In this study the peculiar Andean grass genus Aphanelytrum, with two species, is reduced to Poa sub-
sect. Aphanelytrum comb. & stat. nov. A third species, Festuca reclinata, is assigned to the subsection, 
which shows states transitional between a more typical Poa and Aphanelytrum. Poa subgen. Poa supersect. 
Homalopoa sect. Dioicopoa subsect. Aphanelytrum comb. & stat. nov. is characterized in having stooling 
perennials with decumbent to spreading culm bases that continuously branch and often root at low to 
mid-culm nodes, glabrous spikelets with long rachillas 1.2–4.2 mm long, short glumes less than ½ the 
length of the florets, and lemmas with bifid apexes that are mucronate to short-awned. We provide for the 
three species taxonomic discussions, morphological and anatomical descriptions, keys, illustrations, and 
a list of specimens. Also, we provide two new names, Poa hitchcockiana nom. nov. and Poa sanchez-vegae 
nom. nov., and one new combination, Poa reclinata comb. nov. A new species, Poa auriculata sp. nov. 
from Peru, not thought to be a member of P. subsect. Aphanelytrum, is presented. It is the first in the genus 
with prominent auricles. In addition, we place Poa apiculata in Poa subgen. Poa supersect. Homalopoa sect. 
Dioicopoa subsect. Tovarochloa comb. & stat. nov.
Resumen
En este estudio el género peculiar de gramíneas andinas Aphanelytrum, representado por dos especias, ha 
sido reducido a Poa subsecc. Aphanelytrum comb. & stat. nov. Una tercera especie es incluida en la sub-
sección, Festuca reclinata, esta presenta características transicionales entre las especies más típicas de Poa 
y Aphanelytrum. Poa subgen. Poa supersecc. Homalopoa secc. Dioicopoa subsecc. Aphanelytrum comb. & 
stat. nov. se caracteriza por incluir plantas perennes, stoloniferas, con los culmos decumbentes o extensos, 
continuamente ramificados, a menudo enraizados en los nudos basales hasta los intermedios, las espi-
PhytoKeys 63: 107–125 (2016)
doi: 10.3897/phytokeys.63.8198
http://phytokeys.pensoft.net
Copyright Paul M. Peterson, Robert J. Soreng. This is an open access article distributed under the terms of the Creative Commons Attribution License 
(CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
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Paul M. Peterson & Robert J. Soreng  /  PhytoKeys 63: 107–125 (2016)108
guillas son glabras con rachillas largas de 1.2–4.2 mm de longitud, las glumas más cortas son de la ½ de 
longitud de los flósculos y las lemas con los ápices bífidos son mucronadas hasta breve-aristadas. Nosotros 
proveemos para las tres especias discusiones taxonómicas, descripciones morfológicas y anatómicas, claves, 
ilustraciones y la lista de las muestras usadas. Tambien, proveemos dos nombres nuevos - Poa hitchcockiana 
nom. nov. y Poa sanchez-vegae nom. nov., y una combinación -Poa reclinata comb. nov. Una nueva espe-
cie - Poa auriculata sp. nov. de Perú es presentada. Esta es la especie primera del género con prominentes 
aurículas. En adición, transferimos Poa apiculata a Poa subgen. Poa supersecc. Homalopoa secc. Dioicopoa 
subsecc. Tovarochloa comb. & stat. nov.
Keywords
Anatomy, Aphanelytrum, descriptions, Dioicopoa, Homalopoa, illustrations, key, morphology, taxonomy, 
Tovarochloa
Introduction
The genus Aphanelytrum (Hack.) Hack., first named without a description by Sodiro 
(1889) with a single species, “A. decumbens Hack.” (ex Sodiro, nom. nud.) is based 
on a single collection he made from Ecuador (Chase 1916). Hackel (1897) formally 
described it as a subgenus of Brachyelytrum P. Beauv.; B. subgen. Aphanelytrum Hack., 
including a single new species Brachyelytrum procumbens Hack. Hackel (1902) sub-
sequently recognized Aphanelytrum as a genus with the single species A. procumbens 
(Hack.) Hack. This species is found at mid- to high elevations (2000−4050 m) in hu-
mid to montane forests in the Andes of Bolivia, Colombia, Ecuador, and Peru (Hitch-
cock 1927; Clayton and Renvoize 1986; Jørgensen and Ulloa Ulloa 1994; Renvoize 
1998; La Torre 2002; Soreng et al. 2003; Lægaard 2005; Sánchez Vega et al. 2007). 
Aphanelytrum peruvianum Sánchez Vega, P.M. Peterson, Soreng & Lægaard, a second 
species, was described recently (Sánchez Vega et al. 2007) from Cajamarca, Peru.
Aphanelytrum procumbens is a peculiar grass for having spikelets with minute and un-
veined glumes and two or three florets that are widely-spaced because of the long flexuous 
rachillas that are ½ to ¾ as long as the florets, the lemmas are keeled and 5-veined (Chase 
1916; Hitchcock 1927; Nicora and Rúgolo de Agrasar 1987; Watson and Dallwitz 
1992; Lægaard 2005). Aphanelytrum peruvianum differs from A. procumbens in having 
1–3(–4)-veined glumes (1–2 mm long), narrow leaf blades (0.2–1.2 mm wide), shorter 
culms (14–24 cm long), shorter internodes (3–13 mm long), shorter 3- or 5-veined lem-
mas (2.2–3.5 mm long), and shorter anthers (2–2.9 mm long) [Sánchez Vega et al. 2007].
The placement and evolutionary relationships of Aphanelytrum have been contro-
versial since its inception. It was originally placed in the subfamily “Festuceae” [Festu-
coideae], applied in the broad sense of Bentham (1881), Hackel (1887), and Hitchcock 
(1935). Hackel mistook the upper individual florets for single-flowered spikelets and 
placed Aphanelytrum in Brachyelytrum [B. procumbens] within tribe Agrostideae, subtribe 
Stipinae. Chase (1916) reinterpreted the spikelet morphology, and placed the genus 
between the subtribes Melicinae and Centothecinae, which at that time were considered 
adjacent subtribes of the subfamily Festucoideae, tribe Festuceae. After major realign-
A revision of Poa subsection Aphanelytrum (Poaceae, Pooideae, Poaeae, Poinae)... 109
ments of the classification of the Poaceae (e.g. Clayton and Renvoize 1986), the genus 
was placed in a much more narrowly defined subfamily Pooideae (syn. Festucoideae), 
tribe Poeae (syn. Festuceae) near Poa L. (Clayton and Renvoize 1986). Aphanelytrum 
and Poa have very similar leaf anatomical characteristics, as well as multi-flowered, mem-
branous spikelets with 5-veined, keeled lemmas, glabrous ovaries, and caryopses with 
an oval hilum (Clayton and Renvoize 1986; Clayton et al. 2006). In the most recent 
Poaceae classification (e.g. Soreng et al. 2015) Aphanelytrum is placed as a synonym of 
Poa in the subtribe Poinae, as suggested in studies by Gillespie et al. (2008), Soreng et al. 
(2007, 2015), Soreng and Peterson (2012), and Refulio Rodriguez et al. (2012).
Based on ITS sequences, Gillespie et al. (2008) found the monotypic Tovarochloa 
(T. peruviana T.D. Macfarl. & P. But) to be a weakly supported sister of Aphanelytrum 
procumbens and A. peruvianum. In a plastid trn-TLF derived tree, Refulio Rodriguez et 
al. (2012) verified this result showing a strongly supported Aphanelytrum procumbens 
sister to Tovarochloa, and published a section in Poa for Tovarochloa (nom. inval.; later 
validated as P. sect. Tovarochloa (T.D. Macfarl. & P. But) Molinari, see Molinari-No-
voa 2015), but they left the genus Aphanelytrum unplaced within Poa. Tovarochloa pe-
ruviana is a small, delicate and diminutive annual species with 1-flowered spikelets that 
was previously linked to Dissanthelium Trin. (Clayton and Renvoize 1986; Macfarlane 
and But 1982; Tovar 1993), now both are included within Poa Gillespie et al. 2008; 
Refulio Rodriguez et al. 2012; Soreng et al. 2015). Giussani et al. (in press) estimated 
the time of divergence of Tovarochloa and Aphanelytrum at between 1.24 and 5.05 mya.
Festuca reclinata Swallen, known only from the Páramo del Almorzadero in the 
Cordillera Oriental of Colombia, has been linked to Aphanelytrum based on exhibit-
ing similar morphologies (Stančík and Peterson 2007; Sánchez Vega et al. 2007). The 
growth habit, panicles, and spikelet characteristics of F. reclinata are strikingly similar 
to those found in A. peruvianum and A. procumbens (Sánchez Vega et al. 2007). All 
three species have weak, decumbent culms with intravaginal branching, narrow few-
spikeleted panicles, spikelets with long flexuous rachillas, and small glumes. In com-
parison with F. reclinata, A. peruvianum has smaller lemmas (2.2−3.5 mm vs. 7−8.5 
mm in F. reclinata), smaller anthers (2−2.9 mm vs. 3.5−3.8 mm), and shorter spikelets 
(5−7 mm vs. 10−13 mm) [Sánchez Vega et al. 2007]. The strongly keeled lemmas in 
F. reclinata do not agree with its placement in Festuca (Sánchez Vega et al. 2007). Also 
like Aphanelytrum and Poa, F. reclinata has fused sheath margins, terete rachillas (vs. 
dorsoventrally compressed in Festuca), and lacks the thickened annulate callus typical 
of Festuca. Aphanelytrum procumbens has a caryopsis with short elliptical hilum, less 
than 1/5 the grain in length (typical of Poa vs. linear and proportionally longer in Fes-
tuca), but caryopses have not been observed in A. peruvianum or F. reclinata.
The main goal of this study is to present a systematic revision Poa subsect. Aphan-
elytrum comb. & stat. nov. that includes three species. We make a new combination 
for F. reclinata, and provide new names for Aphanelytrum procumbens and A. peruvi-
anum. In addition, we include a key to the species, complete descriptions, illustrations, 
distribution, specimens examined, and comments for these three species. Furthermore, 
we place Poa apiculata in Poa subsect. Tovarochloa comb. & stat. nov.
Paul M. Peterson & Robert J. Soreng  /  PhytoKeys 63: 107–125 (2016)110
While reviewing Peruvian specimens of Poa, Robert J. Soreng (RJS) found a pe-
culiar collection by John J. Wurdack (Wurdack 1145) from Departamento Amazonas, 
Provincia Chachapoyas located on the summit of Puma-urcu that is similar to Poa 
scabrivaginata Tovar and Festuca reclinata but differed from the forgoing, and all other 
Poa species, in having auriculate collars. We describe this as a new species of Poa, but 
do not include it as a member of P. subsect. Aphanelytrum.
Materials and methods
Herbarium specimens from the following 13 herbaria were examined: AAU, BC, 
COL, CPUN, K, LPB, MA, MO, QCNE, US, USM, USZ, and W (Thiers 2013).
For leaf anatomy, 5 mm long leaf blades were taken from dried herbarium speci-
mens, rehydrated in boiling water, and fixed in FAA for 24 hours. They were trans-
ferred to 70% ethanol, followed by a water rinse and treated for three hours in 50% 
hydrofluoric acid (Martens & Uhl 1980). After being neutralized and washed in water 
the specimens were dehydrated in 2,2-dimethoxypropane (Postek and Tucker 1976) 
and embedded in Polyfin (Polysciences, Inc.) paraffin wax. Transverse serial sections 
were made at 4 μm, stained with buffered Toluidine Blue O (Sakai 1973) and mount-
ed in Lipshaw’s synthetic mounting resin. Photomicrographic images were captured 
using a Zeiss Standard 16WL compound microscope equipped with a Retiga 1300i 
digital camera using ImagePro (MediaCybernetics).
Taxonomic treatment
Poa subsect. Aphanelytrum (Hack.) Soreng & P.M. Peterson, comb. & stat. nov.
urn:lsid:ipni.org:names:77155363-1
within Poa subg. Poa supersect. Homalopoa (Dumort.) Soreng & L.J. Gillespie sect. 
Dioicopoa E. Desv., see Gillespie et al. 2007
Basionym. Brachyelytrum subg. Aphanelytrum Hack., Die Nat. Pflanzenfam., Nachträ-
ge zu Teil II, Abteilung 2. 42. 1897.
Aphanelytrum (Hack.) Hack., Oesterr. Bot. Z. 52: 12. 1902. Aphanelytrum procum-
bens (Hack.) Hack., Oesterr. Bot. Z. 52: 13, text f. 1902.
Aphanelytrum Hack. ex Sodiro, Anales Univ. Centr. Ecuador 3(25): 480. 1889, 
nom. nud.
Type species. Based on Brachyelytrum procumbens Hack. ≡ Aphanelytrum procum-
bens (Hack.) Hack. ≡ Poa hitchcockiana.
Diagnosis. Poa subsect. Aphanelytrum differs from most species of Poa in hav-
ing stooling perennials with decumbent to spreading culm bases that continuously 
branch and often root at low to mid-culm nodes, glabrous spikelets with long rachillas 
A revision of Poa subsection Aphanelytrum (Poaceae, Pooideae, Poaeae, Poinae)... 111
1.2–4.2 mm long, short glumes less than ½ the length of the florets, and lemmas with 
bifid apices that are mucronate to short-awned.
Description. Stooling perennials with intravaginal innovations. Culms 14–80 
(–100) cm tall, decumbent to spreading near base, culm bases that continuously branch 
and often root at low to mid-culm nodes. Leaf blades 3–14 cm long, 0.2–5 (–5.5) 
mm wide, flat to loosely involute; ligules 1–3 mm long, membranous. Panicles few-
flowered with 5–22 spikelets. Spikelets 5–18 mm long, 2–4-flowered, membranous, 
glabrous, disarticulating above the glumes and between the florets; rachilla 1.2–4.2 
mm long, terete in cross section, often prolonged above upper floret; callus glabrous; 
glumes 0.1–3.5 mm long, less than ½ the length of the florets, 0–3 (–4)-veined; lem-
mas 2.2–9.6 mm long, 3- or 5-veined, lanceolate or ovate, apex bifid, mucronate to 
short-awned, if awned up to 2 mm long; paleas 2–7 mm long, apex bifid; lodicules 2, 
glabrous; stamens 3, anthers 2–4.7 mm long; ovaries glabrous. Caryopses compressed 
laterally or unknown.
Three species of northern to central Andes of South America.
Key to the species of Poa subsection Aphanelytrum
1 Glumes veinless, 0.1−0.5 (–0.7) mm long, minute or absent ........................
 ....................................................................................... Poa hitchcockiana
– Glumes veined, 1−3.6 mm long, lower glumes 1 or 3-veined, upper glumes 3 
or 4-veined .................................................................................................2
2 Spikelets 10–13 cm long, 4-flowered; lemmas 6.6–8 mm long; leaf blades 
3–5 mm wide; paleas 4.6–5.2 mm long; culm internodes 14–80 mm long ...
 ............................................................................................... Poa reclinata
– Spikelets 5–7 mm long, 3-flowered; lemmas 2.2–3.5 mm long; leaf blades 
0.2–1.2 mm wide; paleas 2–3.2 mm long; culm internodes 3−18 mm long ..
 ....................................................................................... Poa sanchez-vegae
Poa hitchcockiana Soreng & P.M. Peterson, nom. nov.
urn:lsid:ipni.org:names:77155364-1
Figs 1, 2D, 4C
Brachyelytrum procumbens Hack., Die Nat. Pflanzenfam., Nachträge zu Teil II, Abtei-
lung 2. 42. 1897. Aphanelytrum procumbens (Hack.) Hack., Oesterr. Bot. Z. 52: 
13, text f. 1902.
Aphanelytrum decumbens Hack. ex Sodiro, Anales Univ. Centr. Ecuador 3(25): 480. 
1889, nom. nud.
Type. ECUADOR. Crescit in silvis opacis regionis subandinis, 2000 m, Jul 1887, L. 
Sodiro, s.n. (holotype: W-19813 seen digitally!; isotype: US-865406 fragm. ex W!).
Paul M. Peterson & Robert J. Soreng  /  PhytoKeys 63: 107–125 (2016)112
Figure 1. Poa hitchcockiana: A Habit B Panicle C Sheath, ligule, and blade D Spikelet E Lower floret 
with glumes at base F Glumes at base of lower floret G Lemma H Palea, ventral view I Stamens with 
ovary J Lodicules at base of ovary K Pistil, lodicules at base L Caryopsis M cross section of caryopsis. 
A–D, J–K (Peterson 16571 & Refulio Rodriguez) E, F, L, M (Apollinaire 717 & Arthur, US-913275).
A revision of Poa subsection Aphanelytrum (Poaceae, Pooideae, Poaeae, Poinae)... 113
Description. Straggling and stooling perennials with intravaginal innovations. 
Culms 30–80 (–100) cm tall, decumbent to erect, delicate, glabrous; nodes 3–8 
(–14); internodes 2–14 cm long. Leaf sheaths ½ to 4/5 as long as the internodes, 
membranous to hyaline, often shiny, upper sheaths open ½ the length, keeled; lig-
ules 1–2 mm long, membranous to hyaline, apex erose, often lacerate; blades 6–14 
cm long (flag leaf usually 4–6 cm long), 1.5–4.2 (–5.5) mm wide, flat, thin, lax, 
linear. Panicles 5–18 (–22) cm long, 2–5 cm wide, few-flowered with 10–22 spike-
lets, oblong; branches flexuous, effuse and spreading, the lower branches capillary 
with 2–5 spikelets immediately branching below, the upper branches usually with 
2 spikelets. Spikelets 8–18 mm long, 2- or 3-flowered (often appearing 1-flowered 
with disarticulation of upper florets), laterally compressed, greenish; disarticulation 
between the florets; all florets usually perfect; rachilla joints 1.5–4.2 mm long, often 
prolonged above upper floret; glumes 0.1–0.5 (–0.7) mm long, without veins, min-
ute or absent, apex acute or irregularly lobed or toothed; lemmas 5–9.6 mm long, 
5-veined, lanceolate, apex acuminate, mucronate or short-awned, the awn up to 2 
mm long; paleas 4–7 mm long, 2-keeled, apex bifid; lodicules 0.8–1.1 mm long, 
lanceolate, membranous, glabrous; stamens 3; anthers 2.8–4.7 mm long, yellowish; 
ovaries glabrous with two styles and two stigmas. Caryopses 3.25–4.2 mm long, 
compressed laterally, glabrous, hilum short.
Leaf anatomy. The transverse section leaf anatomy of Poa hitchcockiana is C3, 
XyMS+ with non-radiate, spongy chlorenchyma, without adaxial palisade cells. There 
is a single primary vascular bundle associated with the midrib and a sclerenchyma 
girder 2 or 3-cells thick on the abaxial surface (Fig. 2D). Lateral primary vascular bun-
dles are widely spaced and also have a few abaxial sclerenchyma cells.
Phenology. Flowering year round [?], although no collections made in February, 
May, and September.
Distribution. Poa hitchcockiana is found along the paramo or moist jalca vegeta-
tion between 2000–4025 m of the Cordillera de los Andes in Colombia, Ecuador, 
Bolivia, and Peru.
Conservation status. Since the species is widespread it is of least concern (IUCN 
2010). However, the typical size of populations is undocumented and it has been 
noted by the authors that the grass is sought after by grazers since it is often found 
growing among the protection of perennial shrubs.
Etymology. Since the epithets procumbens and decumbens are blocked in Poa 
by the earlier P. procumbens Curtis and P. decumbens (L.) Scop., we provide a new 
name commemorating the “father of American agrostology,” Albert Spear Hitchcock 
(1865–1935).
Comments. The only wide ranging species of Poa subsect. Aphanelytrum, P. 
hitchcockiana also has the most unusual spikelet morphology with extremely long 
rachillas (1.5–4.2 mm long) and very short [0.1–0.5 (–0.7) mm long] to obscure 
or often absent, unveined glumes. Poa rachillas rarely exceed 1.5 mm, but some 
have spikelets with rachillas up to 2 mm long (e.g. P. sect. Secundae V.L. Marsh ex 
Soreng) P. curtifolia Scribn., P. hartzii Gand., P. stenantha Trin.; (P. sect. Cenisiae 
Paul M. Peterson & Robert J. Soreng  /  PhytoKeys 63: 107–125 (2016)114
Figure 2. Leaf blade transverse sections of Poa sanchez-vegae (A–C) and Poa hitchcockiana (D). Poa 
sanchez-vegae: A Entire leaf blade B Midvein showing a single primary vascular bundle with bulliform 
cells (b indicated by arrows) C Lateral primary vascular bundle with a few associated abaxial sclerenchyma 
fibers (sf) and spongy chlorenchyma (chl). Poa hitchcockiana: D Midvein showing a single primary vascu-
lar bundle with abaxial sclerenchyma fibers (sf) and collapsed chlorenchyma.
A revision of Poa subsection Aphanelytrum (Poaceae, Pooideae, Poaeae, Poinae)... 115
Asch. & Graebn.) P. davisii Bor; (P. supersect. Homalopoa) P. bajaensis Soreng, and 
a few other species in the Homalopoa (H) clade (see Soreng et al. 2010). Four species 
of the related genus Nicoraepoa Soreng & L.J. Gillespie have rachillas up to 3 mm 
long (Soreng & Gillespie 2007). However, there are no known species of Poa with 
unveined glumes.
Specimens examined. BOLIVIA. El Beni: Bella Vista, 26 Dec 1923, A.S. 
Hitchcock 22756 (US). La Paz: Murillo, 2450 m, 7 Apr 1981, S.A. Renvoize 4269, 
T.A. Cope & S.G. Beck (MO); 16°08'S, 68°07'W, 2900 m, 18 Mar 1987, J.C. Solo-
mon 16417 (MO, US); 16°10'S, 68°07'W, 3000 m, 1 Mar 1980, J.C. Solomon 5240 
(MO); 3100m, 16 Mar 1982, T. Feuerer 10719B (MO); 7 Apr 1989, Feuerer 5855C 
(MO). Santa Rosa, 3030 m, 4 Aug 1979, S.G. Beck 1085 (US). Nor Yungus, 3300 
m, O. Buchtien 4268 (US); 3250 m, 3 Apr 1981, S.A. Renvoize 4188 & T.A. Cope (K, 
LPB); Franz Tamayo, 14°43'47"S, 69°04'17"W, 3998 m, 18 Jun 2005, A.F. Fuentes 
8338, R. Hurtado, I. Jiménez, E. Cuevas & R. Cuevas (LPB, MO, USZ); Inquisivi, 
16°48'00"S, 67°16'00"W, 3400–3500 m, 9 Mar 1991, M. Lewis 38263 (MO). CO-
LOMBIA. Boyacá: Nevado de Cocuy, 3750 m, 10 Sep 1938, J. Cuatrecasas 1360 
(US); 4025 m, 7 Oct 1972, A.M. Cleef & P.A. Florschultz 5960 (US). Cauca: 3700 
m, 5 Apr 1985, J.R.I. Wood 4784 (MO). Cundinamarca: Bogota, 10 Aug 1859, 
A. Lindig 1009 (MO, US); Paramo de Chipaque, 3300 m, R. Jaramillo M. 5340 
(COL). Meta: Paramo de Sumapaz, 3700 m, A.M. Cleef 7686 (COL). Tolima: 
16 Dec 1984, J.R.I. Wood 4650 (MO). Camino del Verjon, 3100 m, Jul 1911, 
Apollinaire 717 & Arthur (US-727001), Apollinaire 717 & Arthur (US-913275). J. 
Celestino Mutis 5533 (MA, US). ECUADOR. Cañar: Interandina, 3000 m, 2 Jul 
1950, M. Acosta Solís 16962 (US). Carchi: 00°49'00"N, 77°57'00"W, 3800 m, 10 
Mar 1992, S. Lægaard 101662 (AAU); 00°40'00"N, 77°52'00"W, 3400 m, 1 Nov 
1993, W.A. Palacios 11739 (MO, QCNE). Imbabura: 00°20'00"N, 78°00'00"W, 
3600–3650 m, 7–8 Feb 1992, S. Lægaard 101171 (AAU). Loja: Cajanuma, 04°05'S, 
79°12'W, 2700–3100 m, 5 Mar 1987, I. Grignon 84297 (AAU, MO, US). Napo: 
00°56'00"S, 78°23'00"W, 3600 m, 16–18 Nov 1984, S. Lægaard 53356 (AAU, 
QCA, US). Pichincha: Paso de Huanpango, 3280 m, Jul 1928, G. Firmín 439 
(US); Pedregal, 3400 m, 7 Jul 1944, M. Acosta Solís 8333 (US); Montes Pichinchas, 
3700 m, 21 Jan 1856, W. Jameson s.n. (US), Jameson 168 (US); Pasochoa near Qui-
to, 1890, L. Sodiro s.n. (US); La Campiña, 3000 m, T. Holmgren 649 (US). Tun-
gurahua: Cordillera de Llanganates, 3000 m, 16 Nov 1939, E. Asplund 9737 (US). 
PERU. Cusco: Paucartambo, 2800 m, 17 Mar 2002, P.M. Peterson 16571 & N.F. 
Refulio Rodriguez (US, USM); Calca, 3430 m, 17 Mar 2002, P.M. Peterson 16581 & 
N.F. Refulio Rodriguez (US, USM); Pillco, 17 Apr 1967, C. Vargas C. 19264 (US); 
Quispicanche, 13°35'32.8"S, 70°58'39.9"W, 3097 m, 20 Mar 2007, P.M. Peterson 
20582, R.J. Soreng & K. Romaschenko (US, USM); Moquegua: El Abra, 2000 m, 
Mar 1967, C. Vargas C. 19104 (US). San Martín: Huicungo, 7°58'S, 77°20'W, 
2900–3150 m, 27 Jun 1999, A. Cano s.n. (SI, USM); B. León 3797 (USM).
Paul M. Peterson & Robert J. Soreng  /  PhytoKeys 63: 107–125 (2016)116
Poa reclinata (Swallen) Soreng & P.M. Peterson, comb. nov.
urn:lsid:ipni.org:names:77155361-1
Fig. 3
Festuca reclinata Swallen, Contr. U.S. Natl. Herb. 26(6):254. 1949.
Type. COLOMBIA. Departamento Santander, Cordillera Oriental, Paramo de Almor-
zadero, 3500–3700 m, 20 Jun 1940, J. Cuatrecasas & H. Garcia Barriga 9970 (holo-
type: US-1798714!; isotypes: BC-635144 seen digitally!; COL-34839 seen digitally!).
Description. Stooling perennials forming small tussocks with intravaginal innova-
tions. Culms 30–40 cm tall, decumbent to erect, often weak, spreading to prostrate, 
culm bases continuously branch and often root at low to mid-culm nodes, glabrous; 
nodes 2 or 3 in distal half; internodes 1.4–8.0 cm long. Leaf sheaths about 2/3 as long 
as the internodes, membranous, greenish-white, scabrous, upper sheaths open 1/2 the 
length, collars flared; ligules 1–2.5 mm long, membranous, apex acute, ephemeral; 
blades 5–15 cm × 0.3–0.5 mm, flat, thin, lax, green, abaxially scabrous. Panicles 9–10 
× 2–3 cm, few-flowered with 5–8 spikelets, flexuous, ovate, branched; branches gla-
brous. Spikelets 10–13 mm long, 4-flowered, glabrous; obovate; rachilla 1.2–2.4 mm 
long, minutely scabrous; glumes 1.3–3.6 mm long, membranous, lanceolate, green, 
glabrous, upper margins hairy; lower glumes 1.3–2 mm long, 1-nerved, apex acute; 
upper glumes 3–3.5 mm long, less than ½ as long as the florets, 3-veined, apex acumi-
nate; lemmas 6.6–8 mm long, 5-veined, lanceolate, membranous, green, scabrous, 
apex bifid, two-dentate, awned between the teeth, the awn 1–2 mm long; paleas 4.6–
5.2 mm long, membranous, keels scabrous, apex bifid; lodicules 0.6–0.8 mm long, 
lanceolate; anthers 2.7–3.3 mm long; ovaries glabrous. Caryopses not seen.
Leaf anatomy. The leaf anatomy of Poa reclinata is C3, XyMS+ and the transverse 
sections have many widely spaced vascular bundles with small ribs; sclerenchyma is 
under both abaxial and adaxial epidermis, discontinuous, small, extending to the vas-
cular bundles forming girders; bulliform cells are absent; epidermis is sparsely hairy. 
An anatomical description of P. reclinata is also found in Watson & Dallwitz (1992) 
and Stančík & Peterson (2007, figs 13g, 77c–f).
Phenology. Flowering in July.
Distribution. Poa reclinata is known only from the type locality, the paramo of 
the Colombian Cordillera Oriental, Dept. Santander (Stančík and Peterson 2007).
Conservation status. The species is rare and its conservation status is data defi-
cient (IUCN 2010).
Etymology. The specific epithet is probably in reference to the decumbent, spread-
ing or prostrate culms, a frequent characteristic of the species in this subsection of Poa.
Comments. Stančík and Peterson (2007) mentioned that the spikelets and pani-
cles of Festuca [Poa] reclinata were similar to Aphanelytrum procumbens [Poa hitchcocki-
ana], but the glumes in the former are veined. They provisionally placed F. reclinata in 
F. subg. Subulatae (Tzvelev) E.B. Alexeev sect. Glabricarpae E.B. Alexeev, as suggested 
by Aleexev (1986), along with Festuca caldasii (Kunth) Kunth and F. woodii Stančík. 
A revision of Poa subsection Aphanelytrum (Poaceae, Pooideae, Poaeae, Poinae)... 117
Figure 3. Poa reclinata: A Habit B Sheath, ligule, and blade C Spikelet D Lower floret E Upper glumes 
F Lower glume G Lemma H Palea and rachilla, lateral view I Stamens with palea J Lodicules and with 
pistil. (Cuatrecasas 9970 & Garcua Barriga, US-1798714).
Paul M. Peterson & Robert J. Soreng  /  PhytoKeys 63: 107–125 (2016)118
The latter two species of Festuca have open sheaths, flattened and hairy (hispid?) rachil-
las, and lemmas with rounded keels and annulate calluses, whereas Poa reclinata has 
fused sheaths ½ their length, terete rachillas that are minutely scabrous, and lemmas 
with compressed keels and smooth transitions from callus to lemma. We hope to con-
firm the placement of P. reclinata in P. subsect. Aphanelytrum in future DNA studies.
Poa sanchez-vegae Soreng & P.M. Peterson, nom. nov.
urn:lsid:ipni.org:names:77155370-1
Fig. 2A–C, 4A, B, D–L
Aphanelytrum peruvianum Sánchez Vega, P.M. Peterson, Soreng & Lægaard, J. Bot. 
Res. Inst. Texas 1(2): 842. 2007.
Type. PERU. Departamento Cajamarca, Provincia Cajamarca, Distrito Cajamarca, 
Cerro Akumullca, al SO de Cajamarca, sobre la cima de la ladera occidental del Valle 
de Cajamarca (7°14'15"S Lat, 78°29'24"W Long), 3300 m, 20 Mar 2003, I. Sánchez-
Vega 11781, M. Sánchez-Montoya, R. Cueva R. & J. Montoya (holotype: CPUN!; 
isotypes: AAU!, F!, HAO!, HUT!, LOJA!, MICH!, MO!, SI!, US-3472470!, US-
3686568!, USM!).
Description. Caespitose perennials. Culms 14−24 cm tall, with many culms near 
base, primary and secondary culms appressed, somewhat decumbent near base with 
intravaginal branching, culm bases continuously branch and often root at low to mid-
culm nodes; internodes 3−18 mm long, numerous. Leaf sheaths longer than the inter-
nodes, membranous to hyaline, open to near base to open completely to base, slightly 
keeled; ligules 2−3 mm long, membranous to hyaline, decurrent, apex erose often lac-
erate; blades 3−7 cm long (flag leaf ca 1.6 mm long), 0.2−1.2 mm wide, flat to loosely 
involute, thin, linear, apex naviculate. Panicles 1.7−2.5 cm long, few-flowered with 
5−10 spikelets; branches flexuous, the lower branches with two spikelets, the upper 
branches with single spikelet. Spikelets 5−7 mm long, usually 3-flowered, purplish, 
glabrous, disarticulating above the glumes and between the florets; lower and middle 
florets usually staminate; upper florets usually pistillate; rachilla joints 1.2−2 mm long, 
prolonged above the upper floret; glumes 1−2 mm long, subequal, apex acute, often 
mucronate; lower glume linear, 1-veined; upper glume oblanceolate, 3(4-)-veined, 
often toothed or irregularly lobed minutely bifid; lemmas 2.2−3.5 mm long, 3- or 
5-veined, ovate, apex mucronate with two acute lobes on each side of the mucro, 
the mucro 0.1−0.3 mm long; paleas 2−3.2 mm long, 2-keeled, apex bifid; lodicules 
0.7−0.8 mm long, lanceolate, membranous, glabrous; stamens 3; anthers 2−2.9 mm 
long, yellowish to purplish; ovaries glabrous with two styles and two stigmas. Cary-
opses glabrous.
Leaf anatomy. The transverse section leaf anatomy of Poa sanchez-vegae is C3, 
XyMS+ with non-radiate, spongy chlorenchyma, without adaxial palisade cells (Fig. 
2A–C). There are bulliform cells on the adaxial surface on either side of the midveins 
A revision of Poa subsection Aphanelytrum (Poaceae, Pooideae, Poaeae, Poinae)... 119
Figure 4. Poa sanchez-vegae (A, B, D–L) and Poa hitchcockiana (C). Poa sanchez-vegae: A Habit 
B Sheath, ligule, and blade D Panicle E Spikelet F Glumes, showing lower 1-veined and upper 4-veined 
G Glumes, showing lower 1-veined and upper 3-veined H Lemma I Paleas J Palea, anthers, and rachilla 
K Lodicules and pistil L Pistil. (Sánchez Vega 11781, Sánchez Montoya, Cueva R. & Montoya, US-3472470, 
US-3686568). Poa hitchcockiana: C Sheath, ligule, and blade (Peterson 16571 & Refulio Rodriguez).
Paul M. Peterson & Robert J. Soreng  /  PhytoKeys 63: 107–125 (2016)120
primary vascular bundle without additional sclerenchyma (Fig. 2B). However, there 
are a few abaxial sclerenchyma cells associated with the lateral primary vascular bundles 
(Fig. 2C).
Phenology. Flowering in March.
Distribution. Poa sánchez-vegae is known only from the type locality near the 
western highlands of the Cajamarca Valley and is found on rocky sites associated with 
jalca vegetation (humid alpine grass ecosystems) at 3300 m (Sánchez-Vega et al. 2007).
Conservation status. Poa sanchez-vegae is rare and the conservation status is data 
deficient (IUCN 2010). However, the authors unsuccessfully searched for additional 
material of this species at the type locality on 26 Mar 2008, accompanied by two of the 
original collectors, Isidoro Sánchez-Vega and Juan Montoya. The site had been turned 
into a pine plantation.
Etymology. Since the existing specific epithet was occupied in Poa we provide a 
new name commemorating Isidoro Sánchez Vega, a renowned Peruvian Botanist.
Comments. Based on morphological characters, Isidoro Sánchez-Vega in consul-
tation with Simon Lægaard, first identified the type collection of P. sanchesz-vegae as 
an unknown species of Aphanelytrum. In Sánchez Vega et al. (2007), we described 
this taxon as a new Aphanelytrum, and mention that Festuca reclinata superficially re-
sembled the genus. In addition, P. sanchez-vegae aligned near or on an unusually long 
branch within Poa in preliminary cpDNA and ITS sequence analyses (Gillespie et al. 
2007). Gillespie et al. (2008) included three accessions of Poa hitchockiana (≡ Aphan-
elytrum procumbens) that formed a clade sister to P. sanchez-vegae (≡ A. peruvianum) 
which together was sister to two accessions of P. apiculata (≡ Tovarochloa peruviana). 
These results clearly support our classification of P. sanchez-vegae and P. hitchcockiana 
as members of P. subsect. Aphanelytrum.
Other novelties
Poa subsect. Tovarochloa (Macfarl. & P. But) Soreng & P.M. Peterson, comb. & 
stat. nov.
urn:lsid:ipni.org:names:77155398-1
within Poa subg. Poa supersect. Homalopoa (Dumort.) Soreng & L.J. Gillespie sect. 
Dioicopoa E. Desv., see Gillespie et al. 2007
Basionym. Tovarochloa T.D. Macfarl. & P. But, Brittonia 34(4): 478. 1982.
Type species. Based on Tovarochloa peruviana T.D. Macfarl. & P. But ≡ Poa 
apiculata Refulio.
Comments. For consistency in rank, since Poa apiculata Refulio (≡ Tovarochloa 
peruviana) apparently is sister to the three species in Poa subsect. Aphanelytrum, we 
erect Poa subsect. Tovarochloa.
A revision of Poa subsection Aphanelytrum (Poaceae, Pooideae, Poaeae, Poinae)... 121
Poa auriculata Soreng & P.M. Peterson, sp. nov.
urn:lsid:ipni.org:names:77155360-1
Fig. 5
Type. PERÚ. Departamento Amazonas, Provincia Chachapoyas, summit of Puma-
urcu southeast of Chachapoyas, occasional on dry cliff face, 3100–3200 m, 3 Jul 1962, 
J.J. Wurdack 1145 (holotype: US-2382274!).
Diagnosis. Poa auriculata differs from Poa scabrivaginata Tovar in having 4–6-flowered 
spikelets, a glabrous callus, glumes 2–3 mm long, and lemmas 2.5–3.8 mm long.
Description. Caespitose, annuals or short-lived perennials. Culms 40–72 cm tall, 
erect, scabrous, shiny, often weak; nodes 3–5. Leaf sheaths 2/3 to 4/5 as long the 
internodes, membranous, greenish to stramineous, scabrous, upper sheaths open for 
1/3 the length, keeled, summit with prominent triangular auricles; ligules 3.3–5 mm 
long, membranous to hyaline, apex erose, often split down the center; blades 6–15 cm 
long (flag leaf 3–6 cm long), 3–6 mm wide, flat, thin, lax, linear, scabrous. Panicles 
5–11 cm long, 2.5–5 cm wide, ovate, open; branches flexuous, effuse and spread-
ing with numerous spikelets, scabrous, the lower branches capillary. Spikelets 5–7 
mm long, 4–6-flowered, glabrous, ovate, greenish-yellow tinged with purple; rachilla 
0.4–1.0 mm long; glumes 2–3 mm long, membranous, subequal; lower glumes 2–2.5 
mm long, 1-veined, linear lanceolate, apex acuminate; upper glumes 2.4–3 mm long, 
3-veined, the veins not conspicuous, lanceolate, apex acute; lemmas 2.5–3.8 mm long, 
5-veined, lanceolate, membranous; apex acute, unawned; paleas 2.3–3.7 mm long, 
membranous, 2-keeled, the keels scabrous, apex minutely bifid; lodicules 0.4–0.5 mm 
long, ovate, membranous, glabrous; stamens 3; anthers 1.9–2.1 mm long, yellowish; 
ovaries glabrous with two styles and two stigmas. Caryopses not seen.
Phenology. Flowering in June and July.
Distribution. Poa auriculata is known only from the type locality in Cordillera de 
los Andes of Peru near Chachapoyas between 3100–3200 m growing on a dry cliff face.
Conservation status. The species is rare, but its conservation status is data defi-
cient (IUCN 2010).
Etymology. The specific epithet refers to the triangular auricles that are found on 
the summit of the sheaths, a feature that is unique among species of Poa.
Comments. Initially RJS considered Poa auriculata to be related to species in P. 
subsect. Aphanelytrum. We do not place the new species in P. subsect. Aphanelytrum 
because the spikelets have short rachillas and are 4–6-flowered, the habit is annual to 
short-lived perennials with erect culms, and the lemmas are unawned without mucros. 
However, aside from the auricles, P. auriculata is morphologically consistent with the 
300 or so species that reside within P. subg. Poa supersect. Homalopoa. We hope to 
include a sample of this species in upcoming molecular analyses.
Poa scabrivaginata differs from P. auriculata in having 2-flowered spikelets (4–6 
in P. auriculata), a few cobwebby hairs on the callus (verses glabrous), glumes 3.6–4.3 
mm long (verses 2–3 mm), and lemmas 4.2–4.5 mm long (verses 2.5–3.8 mm) [Tovar 
1965]. Poa aequatoriensis Hack., another species possible to confuse with P. auriculata, 
Paul M. Peterson & Robert J. Soreng  /  PhytoKeys 63: 107–125 (2016)122
Figure 5. Poa auriculata: A Habit B Sheath, auricles, ligule, and blade C Spikelet D Glumes E Lower 
glume F Floret G Lemma H Palea, lateral view I Stamens with palea J Pistil enclosed in palea K Lodicules, 
pistil, and rachilla. (Wurdack 1145, US-2382274)
A revision of Poa subsection Aphanelytrum (Poaceae, Pooideae, Poaeae, Poinae)... 123
differs in having 2- or 3-, rarely 4-flowered spikelets, a few cobwebby hairs on the cal-
lus, and lemmas 3.6–4 mm long (Tovar 1965). Poa aequatoriensis is more wide rang-
ing and has been reported in Colombia, Ecuador, and Peru whereas P. scabrivaginata 
is known from the type (Depto. de Huánuco, Tambo de Vaca, J.F. Macbride 4354 
at US) and one other possible collection (Depto. de Cajamarca, Celendin, I. Sánchez 
Vega 2668 at MO) [Sylvester et al. in press].
Acknowledgements
We thank the Smithsonian Institution’s Restricted Endowment Fund, the Scholarly 
Studies Program, Research Opportunities, Atherton Seidell Foundation, Biodiversity 
Surveys and Inventories Program, Small Grants; the National Geographic Society for 
Research and Exploration (Grant No. 8087-06); Alice R. Tangerini for preparing the 
illustrations and anatomical plate; Stan Yankowski for preparing the anatomical sec-
tions; two anonymous reviewers and Clifford Morden for suggesting improvements to 
the manuscript.
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