A Synopsis of the Genus Stipa (Poaceae) in Middle Asia, Including a Key to Species Identification, an Annotated Checklist, and Phytogeographic Analyses

Abstract

The genus Stipa L. comprises over 150 species, all native to the Old World, where they grow in warm temperate regions throughout Europe, Asia, and North Africa. It is one of the largest genera in the family Poaceae in Middle Asia, where one of its diversity hotspots is located. However, identification of Middle Asian Stipa species is difficult because of the lack of new, comprehensive taxonomic studies including all of the species recorded in the region. We present a critical review of the Mid-Asian representatives of Stipa, together with an identification key and taxonomic listing. We relied on both published and unpublished information for the taxa involved, many of which are poorly known. For each taxon, we present a taxonomic and nomenclatural overview, habitat preferences, distribution, altitudinal range, and additional notes as deemed appropriate. We describe four new nothospecies: S. ×balkanabatica M. Nobis & P. D. Gudkova, S. ×dzungarica M. Nobis, S. ×pseudomacroglossa M. Nobis, S. ×subdrobovii M. Nobis & A. Nowak, one subspecies S. caucasica Schmalh. subsp. nikolai M. Nobis, A. Nobis & A. Nowak, and eight varieties: S. araxensis Grossh. var. mikojanovica M. Nobis, S. caucasica var. fanica M. Nobis, P. D. Gudkova & A. Nowak, S. drobovii (Tzvelev) Czerep. var. jarmica M. Nobis, S. drobovii var. persicorum M. Nobis, S. glareosa P. A. Smirn. var. nemegetica M. Nobis, S. kirghisorum P. A. Smirn. var. balkhashensis M. Nobis & P. D. Gudkova, S. richteriana Kar. & Kir. var. hirtifolia M. Nobis & A. Nowak, and S. ×subdrobovii var. pubescens M. Nobis & A. Nowak. Additionally, 12 new combinations, Achnatherum haussknechtii (Boiss.) M. Nobis, A. mandavillei (Freitag) M. Nobis, A. parviflorum (Desf.) M. Nobis, Neotrinia chitralensis (Bor) M. Nobis, S. badachschanica Roshev. var. pamirica (Roshev.) M. Nobis, S. borysthenica Klokov ex Prokudin var. anomala (P. A. Smirn.) M. Nobis, S. holosericea Trin. var. transcaucasica (Grossh.) M. Nobis, S. kirghisorum P. A. Smirn. var. ikonnikovii (Tzvelev) M. Nobis, S. macroglossa P. A. Smirn. var. kazachstanica (Kotuchov) M. Nobis, S. macroglossa var. kungeica (Golosk.) M. Nobis, S. richteriana var. jagnobica (Ovcz. & Czukav.) M. Nobis & A. Nowak, and S. zalesskii Wilensky var. turcomanica (P. A. Smirn.) M. Nobis are proposed, and the lectotypes for 14 taxa (S. arabica Trin. & Rupr., S. bungeana Trin. ex Bunge, S. caspia K. Koch, S. ×consanguinea Trin. & Rupr., S. effusa Mez, S. ×heptapotamica Golosk., S. jacquemontii Jaub. & Spach., S. kungeica Golosk., S. margelanica P. A. Smirn., S. richteriana, S. rubentiformis P. A. Smirn., S. sareptana A. K. Becker, S. tibetica Mez, and Timouria saposhnikovii Roshev.) are designated. In Middle Asia the genus Stipa comprises 98 taxa, including 72 species, four subspecies, and 22 varieties. Of the 72 species of feather grasses, 23 are of hybrid origin (nothospecies). In Middle Asia, feather grasses can be found at elevations from (0 to)300 to 4500(to 5000) m, but most are montane species. The greatest species richness is observed at altitudes between 1000 and 2500 m. Nineteen species grow above 3000 m, but only nine above 4000 m. The number of taxa (species and subspecies) growing in each country also varies considerably, with the highest noted in Kazakhstan (42), Tajikistan (40), and Kyrgyzstan (35). Of the 76 taxa of Stipa (species and subspecies) recorded in Middle Asia, 41 are confined to the region, with some being known only from a single country or mountain range. Distribution maps of selected species are provided.

Full text

Volume 105
Number 1
2020
Annals
of the
Missouri
Botanical
Garden
R
A SYNOPSIS OF THE GENUS STIPA
(POACEAE) IN MIDDLE ASIA,
INCLUDING A KEY TO SPECIES
IDENTIFICATION, AN
ANNOTATED CHECKLIST, AND
PHYTOGEOGRAPHIC ANALYSES
1
Marcin Nobis,
2,3
* Polina D. Gudkova,
3,4
Arkadiusz Nowak,
5,6
Jakub Sawicki,
7
and
Agnieszka Nobis
2
ABSTRACT
The genus Stipa L. comprises over 150 species, all native to the Old World, where they grow in warm temperate regions throughout
Europe, Asia, and North Africa. It is one of the largest genera in the family Poaceae in Middle Asia, where one of its diversity hotspots is
located. However, identification of Middle Asian Stipa species is difficult because of the lack of new, comprehensive taxonomic studies
including all of the species recorded in the region. We present a critical review of the Mid-Asian representatives of Stipa, together with
an identification key and taxonomic listing. We relied on both published and unpublished information for the taxa involved, many of which
are poorly known. For each taxon, we present a taxonomic and nomenclatural overview, habitat preferences, distribution, altitudinal range,
and additional notes as deemed appropriate. We describe four new nothospecies: S. 3balkanabatica M. Nobis & P. D. Gudkova,
S. 3dzungarica M. Nobis, S. 3pseudomacroglossa M. Nobis, S. 3subdrobovii M. Nobis & A. Nowak, one subspecies S. caucasica
Schmalh. subsp. nikolai M. Nobis, A. Nobis & A. Nowak, and eight varieties: S. araxensi s Grossh.var. mikojanovica M. Nobis, S. caucas ica
var. fanica M. Nobis, P. D. Gudkova & A. Nowak, S. drobovii (Tzvelev) Czerep. var. jarmica M. Nobis, S.drobovii var. persicorum M. Nobis,
S.glareosa P. A. Smirn. var. nemegetica M. Nobis, S. kirghisorumP. A. Smirn. var. balkhashensis M. Nobis & P. D. Gudkova, S. richteriana
1
We would like to express our gratitude to the curators of AA, B, BM, BRNU, COLO, E, FR, FRU, G, GAT, GFW, GOET, IFP,
K, KAS, KFTA, KHOR, KRA, KRAM, KUN, KUZ, L, LE, LECB, M, MO, MSB, MW, NY, P, PE, PR, PRC, TAD, TASH, TK, UPS,
W, WA, WU, and Z for their kind hospitality and assistance during visits and for making specimens of Stipa available on loan. We
would like to thank Robert Soreng and the anonymous reviewer for their valuable comments and improvements to our paper. M. N.
would like to thank Beata Sto˙zek (Krak´ow, Poland) and Vitezslav Plaˇsek (Ostrava, Czech Republic) for their technical assistance in
the preparation of SEM and photography. Financial support for this study came from the National Science Center, Poland (grant no.
2013/09/B/NZ8/03287 and 2018/29/B/NZ9/00313). P. G. received support for her research from the Russian Science Foundation
(research project no. 19-74-10067). Author contributions are as follows. Conception and design: M. N. Field studies, and collection
and determination of plant material: M. N., Ar. N., Ag. N, P. G, and J. S. Revision of herbarium materials: M. N. and P. G. Species
concept, key to species identification, and lectotypifications: M. N. Drafting the manuscript and preparing figures: M. N. with input
of Ag. N. Final approval of manuscript: all authors.
2
Institute of Botany, Faculty of Biology, Jagiellonian University, Krak´ow, Poland.
3
Research Laboratory “Herbarium,”National Research Tomsk State University, Tomsk, Russia.
4
Faculty of Biology, Altai State University, Barnaul, Russia.
5
Botanical Garden-Centre for Biological Diversity Conservation, Polish Academy of Sciences, Warszawa, Poland.
6
Institute of Biology, Opole University, Opole, Poland.
7
Department of Botany and Nature Protection, Faculty of Biology and Biotechnology, University of Warmia and Mazury,
Olsztyn, Poland.
* Author for correspondence: m.nobis@uj.edu.pl, mnobis1577@gmail.com
VERSION OF RECORD FIRST PUBLISHED ONLINE ON 30 APRIL 2020 AHEAD OF SPRING 2020 ISSUE.
doi: 10.3417/2019378 ANN.MISSOURI BOT.GARD. 105: 1–63.

Kar. & Kir. var. hirtifolia M. Nobis & A. Nowak, and S. 3subdrobovii var. pubescens M. Nobis & A. Nowak. Additionally, 12 new
combinations, Achnatherum haussknechtii (Boiss.) M. Nobis, A. mandavillei (Freitag) M. Nobis, A. parviflorum (Desf.) M. Nobis, Neotrinia
chitralensis (Bor) M. Nobis, S. badachschanica Roshev. var. pamirica (Roshev.) M. Nobis, S.borysthenica Klokov ex Prokudin var.
anomala (P. A. Smirn.) M. Nobis, S. holosericea Trin. var. transcaucasica (Grossh.) M. Nobis, S. kirghisorum P. A. Smirn. var. ikonnikovii
(Tzvelev) M. Nobis, S. macroglossa P. A. Smirn. var. kazachstanica (Kotuchov) M. Nobis, S. macroglossa var. kungeica (Golosk.) M. Nobis,
S. richteriana var. jagnobica (Ovcz. & Czukav.) M. Nobis & A. Nowak, and S. zalesskii Wilensky var. turcomanica (P. A. Smirn.) M. Nobis
are proposed, and the lectotypes for 14 taxa (S. arabica Trin. & Rupr., S. bungeana Trin. ex Bunge, S. caspia K. Koch, S. 3consanguinea
Trin. & Rupr., S. effusa Mez, S. 3heptapotamica Golosk., S. jacquemontii Jaub. & Spach., S. kungeica Golosk., S. margelanica P. A.
Smirn., S. richteriana,S. rubentiformis P. A. Smirn., S. sareptana A. K. Becker, S. tibetica Mez, and Timouria saposhnikovii Roshev.) are
designated. In Middle Asia the genus Stipa comprises 98 taxa, including 72 species, four subspecies, and 22 varieties. Of the 72 species of
feather grasses, 23 are of hybrid origin (nothospecies). In Middle Asia, feather grasses can be found at elevations from (0 to)300 to 4500(to
5000) m, but most are montane species. The greatest species richness is observed at altitudesbetween 1000 and 2500 m. Nineteen species
grow above 3000 m, but only nine above 4000 m. The number of taxa (species and subspecies) growing in each country also varies
considerably, with the highest noted in Kazakhstan (42), Tajikistan (40), and Kyrgyzstan (35). Of the 76 taxa of Stipa (species and
subspecies) recorded in Middle Asia, 41 are confined to the region, with some being known only from a single country or mountain range.
Distribution maps of selected species are provided.
Key words: Checklist, distribution, feather grasses, identification key, mountains of central Asia, Old World, Stipa, taxonomy,
typification.
The vascular plant flora of Middle Asia comprises
more than 8000 species, and over 20% of them are
considered endemic (Kamelin, 2002). The mountains of
central Asia have been identified as one of the world’s
biodiversity hotspots (Brooks et al., 2006; Mittermeier
et al., 2011; Critical Ecosystem Partnership Fund,
2018) and one of the 11 most important focal points
of scientific studies and conservation efforts (Giam
et al., 2010). Middle Asia is also characterized by a
high degree of grass diversity, of which one of the
highly represented genera is Stipa L. After the split-
ting of the genus and division of the Eurasian species
into multiple genera (e.g., Achnatherum P. Beauv.,
Aristella Bertol., Celtica F. M. V´azquez & Barkworth,
Macrochloa Kunth, Neotrinia (Tzvelev) M. Nobis,
P. D. Gudkova & A. Nowak, Oloptum R¨oser & Hamasha,
Orthoraphium Nees, Ptilagrostis Griseb., Patis Ohwi,
Piptatherum P. Beauv., Psammochloa Hitchc., Stipellula
R¨oser & Hamasha, and Trikeraia Bor), Stipa s. str. still
comprises more than 150 species that are distributed in
warm temperate regions throughout Europe, Asia, and
North Africa (Roshevitz, 1934; Bor, 1960,1970; Tzvelev,
1968, 1976; Martinovsk´y, 1980; Freitag, 1985; Jacobs &
Everett, 1996; Kotukhov, 2002; Wu & Phillips, 2006;
Barkworth, 2007; Barkworth et al., 2008; Romaschenko
et al., 2008, 2012; Nobis, 2013, 2014; Hamasha et al.,
2012). Its representatives grow in open grasslands, alpine
meadows (in neutral to alkaline soils), lowlands, moun-
tains, and stony and sandy deserts, as well as on steppes,
rocks, and screes. Sometimes particular Stipa species
create communities, commonly known as feather grass
steppes or feather grass grasslands, in which they con-
stitute diagnostic, constant, and/or dominant species
(e.g., Koelerio macranthae–Stipetum pennatae Kolbek,
Stipetum tirsae Meusel, Festuco valesiacae–Stipetum
capillatae Sillinger, Asperulo albiflorae–Stipetum zerav-
shanicae M. Nobis, A. Nowak & A. Nobis, or Stipetum
lipskyi A. Nowak, S. Nowak, A. Nobis & M. Nobis;
Korovin, 1962; Stanyukovich, 1982, Lavrenko et al.,
1991; Gadghiev et al., 2002; Agakhanjanz & Breckle,
2003; Eberhardt, 2004; Peer et al., 2008; Chytr´y, 2010;
Ermakov, 2012; Rachkovskaya & Bragina, 2012; Kabaˇs
et al., 2013; Nobis et al., 2013; Nowak et al., 2016, 2018).
Stipa comprises perennial and usually densely tufted
plants whose most distinctive character is a terminal
awn on the lemma. The awn can be unigeniculate or
bigeniculate, scabrous/glabrous to variously pilose, and
can reach from 2 to 50 cm in length. Stipa clearly differs
from the other Old World genera of Stipeae in the
pattern of the lemma micromorphology, which has great
phylogenetic value within the tribe. Stipa species have
lemmas characterized by the presence of rectangular to
square fundamental (long) cells, with numerous hooks
and sparse, reniform to oblong silica bodies sometimes
associated with cork cells. Hooks are distributed on
short cells along the entire upper surface of the lemma
(Barkworth & Everett, 1987; Romaschenko et al., 2012;
Nobis, 2013; Nobis et al., 2016a, 2019a, 2019b;
Olonova et al., 2016). This pattern of lemma epidermis
has been called saw-like, in contrast to maize-like, which
is characterized by short fundamental cells and extremely
numerous silica bodies on the upper surface of the
lemma, typical for achnatheroid grasses represented by
e.g., Achnatherum,Aristella,orStipellula (Romaschenko
et al., 2012). The third type of lemma epidermis pattern
encountered in Stipeae and observed in Neotrinia,
Orthoraphium,Piptatherum,Psammochloa,Ptilagrostis,
and Trikeraia is characterized by elongate fundamental
(long) cells that vary in shape, from rectangular to elon-
gated with straight or sinuous edges, more or less nu-
merous silica bodies associated with cork cells, and
sparse or absent hooks (Romaschenko et al., 2012,
2014; Nobis & Nobis, 2013; Nobis et al., 2019a,
2019b). Some authors have proved that although the
2 Annals of the
Missouri Botanical Garden

general pattern of the lemma epidermis in speciesof Stipa
is relatively uniform, it may be useful in the identification
of particular species within and/or between different
generic sections (Nobis, 2013, 2014; Nobis et al.,
2014b, 2015b, 2016a; Olonova et al., 2016). Despite
several phylogenetic studies on Stipeae carried out so far
(Romaschenko et al., 2008, 2010, 2012; Cialdella et al.,
2010; Hamasha et al., 2012; Sclovich et al., 2015), no
final phylogenetic relationships have been identified within
the genus Stipa. In previous studies, based on analysis of ITS
and various regions of chloroplast DNA, phylogenetic res-
olution between species was obscure; when clades were
resolved, support was often weak (Hamasha et al., 2012;
Romaschenko et al., 2012). Recently, Krawczyk et al.
(2017) showed that the nuclear ribosomal RNA intergenic
spacer (IGS) region, especially the part adjacent to the 26S
nuclear ribosomal DNA, can serve as an effective molecular
marker enabling reconstruction of the phylogeny of Stipa.
Using this region, the authors conducted a molecular phy-
logenetic study for 36 species of Stipa, obtaining the first
phylogenetic tree, with well-resolved clades, linked in most
cases to infrageneric sections, which are distinguished by
morphological characters. Nevertheless, as of yet no
comprehensive explanation of phylogenetic relationships
within the genus Stipa has been put forward.
Because of their ornamental value, feather grasses
have drawn the attention of nature lovers as well as
botanists for many years. The first taxonomic studies
including Stipa from Middle Asia (the region including
southern Kazakhstan, western China, Kyrgyzstan, Taji-
kistan, Uzbekistan, Turkmenistan, northern Afghani-
stan, northwestern Iran, and northern Pakistan) were
published in the 19th century (Trinius, 1820, 1829;
Link, 1827; Trinius & Ruprecht, 1842; Grisebach,
1852, 1868; Steudel, 1855; Hackel, 1887; Hooker,
1896). At the beginning of the 20th century, research
on Stipa was conducted by Junge (1910), Roshevitz
(1916, 1920, 1929, 1934), Smirnov (1924, 1925, 1928,
1929a, 1929b, 1930, 1934), and Drobov (1941). After
World War II, data on the taxonomy of the genus Stipa
were supplemented by Gamayunova (1956), Ovchinnikov
and Chukavina (1957), Bor (1960, 1970), Pazij (1968),
Tzvelev (1968, 1974, 1976), Martinovsk´y (1980), Cope
(1982), Freitag (1985), and Kuo and Sun (1987). The
number of Stipa species recognized for the region varied
between 40 and 70. However, these numbers also in-
cluded several taxa (based on morphological and mo-
lecular studies) currently belonging to other genera,
e.g., Ptilagrostis,Achnatherum,orStipellula.Onthe
other hand, during the last 30 years, some new species
of Stipa also have been described from central Asia by
Lomonosova (1990), Kotukhov (1987, 1991, 1994, 1998a,
1998b), Tzvelev (2012), Nobis (2013, 2014), Nobis et al.
(2013, 2014b, 2017a), and Zhao and Guo (2011, 2017).
Further exploration of the region is expected to reveal
additional species. Approximately half of the Middle Asian
feather grasses are endemics (restricted either to a single
country or mountain range, or to the region in general) and
are considered threatened (Pazij, 1968; Tzvelev, 1976;
Nobis, 2009). It is worth mentioning that the system-
atic position of several taxa representing the genus is
still unclear and requires further study.
Based on morphological comparison, some researchers
have hypothesized that many Stipa taxa have arisen
through hybridization (Smirnov, 1970; Tzvelev, 1976;
Kotukhov, 2002; Nobis, 2013). Nevertheless, before
now, all information and hypotheses on the hybrid
origin of these taxa, as well as on hybridization events
in feather grasses, were based on morphology only.
Recently Nobis et al. (2019c) provided the first mo-
lecular evidence for this phenomenon in Stipa.Hy-
bridization within this genus is particularly interesting
because hybrids are perennial and reproduce vegeta-
tively and, less frequently, sexually (Nobis, 2013;
Nobis et al., 2017a). Most produce fertile pollen grains
and therefore are, or may be, able to backcross with
parental species, resulting in introgression (Nobis
et al., 2019c). We presume that currently more than
30% of Stipa species have a hybrid origin.
An additional impediment to the identification of
Stipa in the region is the lack of current comprehensive
treatment of taxa in the region. Thus, the main goals of
this paper are to provide, for members of Stipa currently
known from the region: (1) an identification key; (2) a
taxonomic and nomenclatural summary that also in-
cludes notes on each taxon’s habitat preferences and
distribution; (3) descriptions of new taxa found by our
research team during field studies or revision of her-
barium materials; and (4) an analysis of the diversity
and distribution patterns of Stipa in Middle Asia.
MATERIALS AND METHODS
This study is based on the authors’fieldwork in the
region (from 2007 to the present) and examination of
specimens deposited in the following herbaria: AA, B,
BM, BRNU, COLO, E, FR, FRU, G, GAT, GFW,
GOET, IFP, K, KAS, KFTA, KHOR, KRA, KRAM,
KUN, KUZ, L, LE, LECB, M, MHA, MO, MSB, MW,
NY, P, PE, PR, PRC, TAD, TASH, TK, UPS, W, WA,
WU, and Z as well as critical review of available liter-
ature regarding the taxonomy and distribution of feather
grasses. Herbarium codes mentioned above follow
Thiers (2018). In total, more than 5000 sheets with
specimens representing the genus Stipa were revised. The
specimens were collected from Middle Asia (Kamelin,
1973) as well as the mountains of central Asia. This area
is located in the western part of central Asia and
comprises the following countries: southern Kazakhstan,
Kyrgyzstan, Tajikistan, Uzbekistan, Turkmenistan, northern
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Nobis et al. 3
Synopsis of Stipa (Poaceae) in Middle Asia

Afghanistan, northwestern Iran, and northern Pakistan
(Fig. 1). The most important morphological characters
used in the taxonomy of Stipa are presented in Figure 2.
Illustrations and descriptions of other morphological
characters can be found, e.g., in the work of Roshevitz
(1934), Martinovsk´y (1967, 1970, 1972, 1976, 1977),
Freitag (1985), Moraldo (1986), Gonzalo et al. (2012,
2013), Nobis (2013, 2014), Gudkova et al. (2013b),
and Nobis et al. (2013, 2016a, 2017a).
MORPHOLOGICAL ANALYSES
The numerical analyses are based on 76 taxa (species
and subspecies) representing the genus Stipa in Middle
Asia. We selected 16 variable morphological characters
(Table 1) and scored all taxa of Stipa examined, as well
as 22 additional taxa selected from six genera of the
tribe Stipeae. Each taxon was treated as an operational
taxonomic unit (OTU) in accordance with the methods
used in numerical taxonomy (Sokal & Sneath, 1963).
Cluster analysis was performed on all OTUs to estimate
morphological similarities among the species. The sim-
ilarities among OTUs were calculated using Gower’s
general similarity coefficient. A cluster analysis (un-
weighted pair group method with arithmetic mean) was
carried out using PAST software (Hammer et al., 2001).
TAXONOMIC TREATMENT
Within the the key and list of taxa presented below,
four new nothospecies (Stipa 3balkanabatica M. Nobis
& P. Gudkova, S. 3dzungarica M. Nobis, S. 3pseudo-
macroglossa M. Nobis, and S. 3subdrobovii M. Nobis &
A. Nowak), one subspecies (S. caucasica Schmalh. subsp.
nikolai M. Nobis, A. Nobis & A. Nowak), and eight
varieties (S. araxensis Grossh. var. mikojanovica M. Nobis,
S. caucasica var. fanica M. Nobis, P. D. Gudkova &
A. Nowak, S. drobovii (Tzvelev) Czerep. var. jarmica
M. Nobis, S.drobovii var. persicorum M. Nobis, S.glareosa
P. A. Smirn. var. nemegetica M. Nobis, S. kirghisorum
P. A. Smirn. var. balkhashensis M. Nobis & P. D.
Gudkova, S. richteriana Kar. & Kir. var. hirtifolia M.
Nobis & A. Nowak, S. 3subdrobovii M. Nobis & A. Nowak
var. pubescens M. Nobis & A. Nowak) are described. Ad-
ditionally, 12 new combinations, Achnatherum haussknech-
tii (Boiss.) M. Nobis, A. mandavillei (Freitag) M. Nobis,
A. parviflorum (Desf.) M. Nobis, Neotrinia chitralensis (Bor)
M. Nobis, S. badachschanica Roshev. var. pamirica
(Roshev.) M. Nobis, S.borysthenica Klokov ex Prokudin
var. anomala (P.A.Smirn.)M.Nobis,S. holosericea Trin.
var. transcaucasica (Grossh.) M. Nobis, S. kirghisorum P. A.
Smirn. var. ikonnikovii (Tzvelev) M. Nobis, S. macroglossa
P. A. Smirn. var. kazachstanica (Kotuchov) M. Nobis,
S. macroglossa var. kungeica (Golosk.) M. Nobis, S. rich-
teriana var. jagnobica (Ovcz. & Czukav.) M. Nobis &
A. Nowak, S. zalesskii Wilensky var. turcomanica (P. A.
Smirn.) M. Nobis, are proposed, and the lectotypes for 14
taxa, namely S. arabica Trin. & Rupr., S. bungeana Trin.
ex Bunge, S. caspia K. Koch, S. 3consanguinea Trin. &
Rupr., S. effusa Mez, S. 3heptapotamica Golosk.,
S. jacquemontii Jaub. & Spach., S. kungeica Golosk.,
Figure 1. The area of study.
4 Annals of the
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Figure 2. Selectedmorphologicalcharacters offeather grasses. —A. Awn and anthecium: width of awn column (AC), length of anthecium
(AL), length of callus (CalL), length of hairs on the ventral part of the callus (CvH), length of hairs on the dorsal part of the callus (CdH), length of
the lower segment of column (Col
1
L), length of the upper segment of column (Col
2
L), length of the peripheral ring of the callus base (CRL),
width of the peripheral ring of the callus base (CRW), distance from the end of the dorsal line of hairs to the top of the lemma (DDL), distance
from the end of the ventral line of hairs to the top of the lemma (DVL), length of hairs in the dorsal line on the lemma (LHD), length of hairs in the
ventral line on the lemma (LHV), length of hairs on the seta (SHL), length of the longest hairs on the seta (SL). —B. Top of the lemma (from left):
with well-developed ring of hairs, with poorly developed ring of hairs, and without ring of hairs. —C. Micromorphological patterns of the lemma
epidermis. —D. Adaxial (upper) surface of leaves (from left): with mixed short and long hairs, with hairs up to 2 mm long, and with dense hairs
up to 6 mm long. —E. Ring (foot) of the callus (from left): pyriform, cuneate, goblet-like concave and ovate in shape. —F. Anthecium. tl, top of
lemma; d, dorsal line of hairs; sd, subdorsal line of hairs; sv, subventral line of hairs; v, ventral line of hairs; fc, foot of the callus.
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Nobis et al. 5
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S. margelanica P. A. Smirn., S. richteriana,S. rubentiformis
P. A. Smirn., S. sareptana A. K. Becker, S. tibetica Mez,
and Timouria saposhnikovii Roshev., are designated. For
seven species (S. 3adamii M. Nobis, S. 3albasiensis L. Q.
Zhao & K. Guo, S. 3assyriaca Hand.-Mazz., S. 3consan-
guinea,S. 3gnezdilloi Pazij, S.3heptapotamica,and
S. 3pseudocapillata), we indicate a status of nothospecies
(pro species). In addition, for all taxa with a hybrid origin
presented in the list below, we provide information on their
putative parental species.
For each examined taxon, we provide the following
information: type specimen (holotype or lectotype)
and place of its preservation, habitat require-
ments, geographic distribution, altitudinal range,
and other chorological, taxonomic, or nomenclatural
information. Taxonomic synonyms for species given
below are based on the revision of original herbar-
ium material of the species conducted by the two
firstauthorsofthepaperaswellastheworksof
Tzvelev (1976), Freitag (1985), Wu and Phillips
(2006), Nobis (2010, 2013, 2014), Nobis and
Gudkova (2011, 2016), Nobis et al. (2015b,
2016b), Nobis and Klichowska (2016), and working
lists of plant species available online through Plant
List (,http://www.theplantlist.org/.) and Tropicos®
(,www.tropicos.org.).
CHECKLIST OF STIPA
Stipa L., Sp. Pl. 1: 78. 1753. TYPE: Stipa pennata L.
KEY TO TAXA OF STIPA IN MIDDLE ASIA
1. Awn scabrous throughout, covered with hairs up to 0.3 mm long (shorter than diameter of the awn) . . .............2
19. Awn entirely or only partially covered with hairs over 0.3 mm long (longer than diameter of the awn) . . . . . . . . . . . . . 8
2. Glumes 9–15 mm long, callus 0.5–1.3 mm long, anthecium 5–7 mm long . . ...................................3
29. Glumes 20–35 mm long, callus 1.5–4 mm long, anthecium 7–14 mm long . . ...................................4
3. Callus 1–1.3 mm long, awn with hairs up to 0.1 mm long, ligules of vegetative shoots 0.2–0.5 mm long, lemma with 7
linesofhairs,ofwhichthe dorsaloneterminatesbelowthehalfof thelemmalength........S. bungeana Trin. ex Bunge
39. Callus (0.5–)0.6–0.8(–1) mm long, awn with hairs 0.2–0.3 mm long, ligules of vegetative shoots up to 0.2 mm long,
densely hairy on margins, lemma with indistinct lines of hairs or pilose all around, hairs terminate in the upper half of
the lemma length . . . ........................ S. richteriana Kar. & Kir. subsp. jagnobica (Ovcz. & Czukav.) Tzvelev
4. Abaxial surface of blades of vegetative leaves usually glabrous, rarely slightly scabrous, adaxial surface densely covered
with hairs up to 0.1 mm long, anthecium with well-developed ring of hairs at the apex . . .........................5
49. Abaxial surface of blades of vegetative leaves usually scabrous, rarely almost glabrous, adaxial surface densely covered
with hairs 0.2–0.5 mm long, or a mixture of short and long hairs, anthecium without or with poorly developed ring of
hairs at the apex . . ...................................................................................6
5. Ligules of vegetative leaves up to 0.2 mm long, callus 2.1–4.1 mm long . . .....................S. krylovii Roshev.
59. Ligules of vegetative leaves 0.3–2 mm long, callus 1.5–2 mm long . . . . .................S. margelanica P. A. Smirn.
6. Lemma with poorly developed ring of hairs at the apex (rarely well developed), abaxial surface of leaves of vegetative
shoots scabrous due to prickles and spinules and adaxially covered with short or mix of short and long hairs (long
hairspresentonly onmarginal ribs).............................................S. sareptana A. K. Becker
69. Lemma glabrous at the apex or rarely with scattered hairs and/or prickles near the lemma margins, abaxial surface of
leaves of vegetative shoots scabrous to almost glabrous, and densely covered adaxially with 0.2–0.5 mm long hairs
(rarely, long hairs are present only on marginal ribs) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7. Anthecium 7–11 mm long, awn 7–11 cm long, internodes usually longer than culm sheaths . . . S. karakabinica Kotukhov
79. Anthecium (10–)11–13(–14) mm long, awn (10–)12–22 cm long, internodes usually shorter than culm sheaths . . .
......................................................................................... S. capillata L.
8(1). Column (lower segment of awn) plumose with hairs distinctly longer than those on seta (upper segment of awn) . . . . . . . . . 9
89. Column glabrous, scabrous or plumose with hairs equal to or shorter than those on seta . . . . .....................14
9. Awn with hairs on seta 0.1–0.3 mm long, 5 or more times shorter than those on column . . . . . . . . . . . . . . . . . . . . . . . 10
99. Awn with hairs on seta (0.3–)0.5–1.1(–1.4)mm long, 1.5–4 times shorterthan those onthe column . . . S. roborowskyi Roshev.
10. Awn 1.5–3.5 cm long, anthecium 5–8 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
109. Awn 12–23 cm long, anthecium over 11 mm long . . . . . . . . . . . . . . . . . . . . . ......................S. holosericea Trin.
11. Column with hairs up to 1.5 mm long . . . ..................................................S. regeliana Hack.
119. Column with hairs 2–4 mm long . . .....................................................................12
12. Panicle lax, branches more than 3 cm long, anthecium 5–8 mm long . . .................S. penicillata Hand.-Mazz.
129. Panicle contracted, branches up to 3 cm long, anthecium 4–6 mm long . ...................................... 13
13. Glumes 6–8(–8.5) mm long, almost equal, abruptly contracted into more or less acute tip, awn usually more than 21 mm
long .............................................................................S. subsessiliflora Roshev.
139. Glumes (9–)9.5–13.5 mm long, distinctly unequal, gradually narrowed into acute tip, awn usually up to 19 mm long . . .
......................................................................... S.basiplumosa Munro ex Hook. f.
14(8). Awn unigeniculate or indistinctly bigeniculate . . .......................................................15
149. Awn distinctly bigeniculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
15. Column glabrous, smooth or scabrous, seta with hairs 3–11.5 mm long . . .....................................16
159. Awn pilose throughout, column with hairs (0.2–)0.3–2.5(–3.5) mm long, seta with hairs (0.3–)0.5–11.5 mm long . . . 29
16. Lemma with 3 longitudinal lines of hairs .................................................................17
169. Lemma with (5)7 longitudinal lines of hairs . . ...........................................................18
6 Annals of the
Missouri Botanical Garden

17. Glumes 28–36(–39) mm long, anthecium (6.9–)8–9.5(–9.8) mm long, callus 0.7–1(–1.2) mm long, glabrous or rarely
with solitary hairs on the ventral part, callus base footlike, expanded with peripheral ring (0.33–)0.35–0.43(–0.45) mm
in diam. . . . . . . ................................................................... S. karataviensis Roshev.
179. Glumes (37–)39–45 mm long, anthecium (9.6–)10–11.5 mm long, callus (1.3–)1.4–1.7 mm long, densely pilose, callus
base not expanded with peripheral ring 0.32–0.37 mm in diam. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. 3adamii M. Nobis
18. Ligules of vegetative shoots up to 0.2 mm long . . . . .......................................................19
189. Ligules of vegetative shoots 0.2–2.1 mm long .............................................................22
19. Leaves of vegetative shoots distinctly scabrous, awn column glabrous and smooth, 37–58 mm long, seta with hairs
8.3–10.5(–11.5) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. longiplumosa Roshev.
199. Leaves of vegetative shoots glabrous and smooth, awncolumn more or less scabrous, (11–)14–28(–32) mm long, seta with
hairs (5–)6–8.5(–9)mm long.............................................................................. 20
20. Seta straight, (5.2–)6–9.6(–12.1) times longer than column, column more or less scabrous due to short prickles, callus
base distinctly footlike, expanded with peripheral ring (0.44–)0.48–0.65(–0.75) mm in diam. ........S. lipskyi Roshev.
209. Seta arcuate or slightly flexuous, rarely straight, (2.6–)3–4.5(–5.1) times longer than column, column distinctly
scabrous due to hard, hooked prickles, callus base not expanded with peripheral ring 0.26–0.42 mm in diam. . . . . . . 21
21. Callus 1.7–2.2(–2.4) mm long, on the ventral surface densely and long pilose, on the dorsal surface with straight
0.5–0.9(–1.1) mm long hairs occurring only in the middle part of the callus (at 0.35–0.50 mm below the base of lemma
glabrous), adaxial surface of leaves of vegetative shoots densely short-pilose with hairs 0.1–0.2(–0.3) mm long . . .
............................................................................ S. 3tadzhikistanica M. Nobis
219. Callus 0.9–1.4(–1.6) mm long, on the ventral surface densely and long pilose, on the dorsal surface with falcate or
straight (1.2–)1.5–2.2 mm long hairs occurring just below the base of lemma, adaxial surface of leaves of vegetative
shoots densely long-pilose with hairs 0.35–0.65 mm long . . . . . .......................... S. 3brevicallosa M. Nobis
22(18). Glumes 24–34 mm long, anthecium (6–)7–9(–9.5) mm long, awn with column (12–)14–19(–22) mm
long.............................................................................S. tianschanica Roshev.
229. Glumes (32–)35–68 mm long, anthecium (10.3–)11–15.5 mm long, awn with column more than 22 mm long . . . . . . . 23
23. Seta with hairs (2.8–)3–4 mm long . . .................................................S. 3brozhiana M. Nobis
239. Seta with hairs (4.2–)4.5–6.7(–8) mm long . . .............................................................24
24. Glumes (32–)35–45(–49) mm long, awn (9.5–)11–16.2(–17.7) cm long, seta (7–)8.5–12(–13.4) cm long . . .........25
249. Glumes 44–68 mm long, awn (15–)17–25 cm long, seta (12–)13–21 cm long . . . . . ............................26
25. Awn (9.5–)11–14(–14.5) cm long, column (2.3–)2.5–3.1 cm long and distinctly scabrous, seta with hairs (6.5–)7–
8(–8.2)mm long......................................................................S. 3hissarica M. Nobis
259. Awn (10.8–)12–16.2(–17.7) cm long, column (2.7–)3–4.6(–5.4) cm long and somewhat scabrous, seta with hairs
(4.2–)4.5–6(–6.3) mm long .................................................................S. 3alaica Pazij
26. Lower segment of awn (column) glabrous and smooth . . .........................S. 3pseudomacroglossa M. Nobis
269. Lower segment of awn (column) scabrous . . .............................................................27
27. Leaves of vegetative shoots distinctly scabrous, glumes 60–68 mm long, awn 21.5–25 cm long...........S. okmirii Dengub.
279. Leaves of vegetative shoots slightly scabrous to almost glabrous, glumes 35–56 mm long, awn 12–20.6(–22.7) cm
long ................................................................................................28
28. Glumes 35–45(–49) mm long, awn 12–16.5(–18.5) cm long, adaxial surface of leaves covered with hairs up to
0.1 mmlong................................................................S. 3manrakica Kotukhov
289. Glumes 48–56 mm long, awn (15.8–)16.7–20.6(–22.7) cm long, adaxial surface of leaves covered with a mix of short
and long hairs up to 0.25 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. 3talassica Pazij
29(15). Seta with hairs up to 1 mm long .....................................................................30
299. Seta with hairs more than 3 mm long . . .................................................................31
30. Awns 11–15 cm long, glumes 32–40 mm long, vegetative leaves distinctly scabrous due to densely distributed spinules
and prickles ...................................................S. 3balkanabatica M. Nobis & P. D. Gudkova
309.Awns7–11 cm long, glumes 24–30 mm long, vegetative leaves glabrous or slightly scabrous due to scattered
prickles................................................................S. 3consanguinea Trin. & Rupr.
31. Seta straight, (5.5–)6.5–12(–14) times longer than column . . ...............................................32
319. Seta falcate, arcuate, flexuous or straight, (1.7–)2.3–4.2(–5) times longer than column . . .......................36
32. Glumes 35–45 mm long, anthecium 9.5–11.2(–11.5) mm long, seta with hairs (3–)3.8–5.5(–6) mm long . . .
........................................................................... S. aktauensis Roshev.
329.Glumes(41–)44–90(–96) mm long, anthecium (11.5–)12–16(–16.5) mm long, seta with hairs (5.5–)6–9.8(–11.5) mm long . . .
........................................................................................................ 33
33. Leaves scabrous abaxially, callus base cuneate (not expanded), lemma without or with poorly developed ring of unequal
hairs at the apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. narynica M. Nobis
339. Leaves glabrous and smooth abaxially, callus base expanded, lemma with well-developed ring of long hairs at the
apex ................................................................................................34
34. Leaves of vegetative shoots flat or convolute, at the adaxial surface densely and very shortly pilose, with hairs
0.05–0.15 mm long, ligules of the vegetative shoots very densely pilose with hairs up to 3 mm long, base of the callus
pyriform . . . . . ......................................................................S. magnifica A. Junge
349. Leaves of vegetative shoots convolute, at the adaxial surface densely and 0.20–0.55 mm long, pilose, ligules of the
vegetative shoots pilose with up to 2-mm-long hairs, base of the callus ovate or broadly cuneate . . ................35
35. Callus densely pilose all over, base of the callus flat and broadly cuneate, lower segment of awn densely pilose . . .
................................................................................. S. ovczinnikovii Roshev.
359. Dorsal part of callus glabrous or shortly pilose only at base of lemma, ventral part of callus more or less densely pilose,
base of the callus goblet-like, concave and ovate in shape, lower segment of awn sparsely pilose . . . S. lingua A. Junge
Volume 105, Number 1
2020
Nobis et al. 7
Synopsis of Stipa (Poaceae) in Middle Asia

36(31). Callus 0.8–1.2(–1.5) mm long, hairs in the dorsal part of the callus strongly falcate . . . . . S. drobovii (Tzvelev) Czerep.
369. Callus (1.6–)1.8–3.5(–4) mm long, hairs in the dorsal part of the callus straight or flexuous . . . . . . . . . . . . . . . . . . . . . 37
37. The longest ligules of the vegetative shoots 0.3–2.5 mm long, if ligules shorter (up to 0.2 mm) than glumes, 18–28 mm
long ................................................................................................38
379. The longest ligules of the vegetative shoots up to 0.2 mm long, glumes 34–61 mm long . . . . . . . . . . . . . . . . . . . . . . . . . 42
38. Glumes (50–)55–61 mm long, awn (10.9–)13.2–16.4(–17.6) cm long, leaves of vegetative shoots densely
pubescent ................................................................S. 3fallax M. Nobis & A. Nowak
389. Glumes up to 45 mm long, awn 4–13.5 cm long, leaves of vegetative shoots glabrous or scabrous, rarely densely
pubescent ...........................................................................................39
39. Awn distinctly unigeniculate, the longest ligules up to 0.3(–0.6) mm long . . . . . . . . . . . . . . . . . S. glareosa P. A. Smirn.
399. Awn indistinctly bigeniculate, the longest ligules 0.8–2.3 mm long . . . ........................................40
40. Awn 11.5–14 cm long, glumes 35–43 mm long, leaves of vegetative shoots glabrous . . ...........S. 3gnezdilloi Pazij
409. Awn up to 10 cm long, glumes 18–38 mm long, leaves of vegetative shoots scabrous . . .........................41
41. Glumes 18–26 mm long, column with hairs 0.5–1 mm long, leaves scabrous to almost slightly scabrous . . .......
..................................................................... S. 3albasiensis L. Q. Zhao & K. Guo
419. Glumes 26–38 mm long, column with hairs 0.3–0.6 mm long, leaves glabrous to slightly scabrous . . . S. 3tzvelevii Ikonn.
42(37). Adaxial surface of leaves of vegetative shoots covered with hairs 0.15–0.6 mm long . . .......................43
429. Adaxial surface of leaves of vegetative shoots covered with prickly hairs 0.08–0.1 mm long . . ...................44
43. Dorsal part of the callus with hairs 1.5–2 mm long and flexuous, callus (1.5–)1.6–2.1(–2.3) mm long . . . . . . . ....
.................................................................... S. 3subdrobovii M. Nobis & A. Nowak
439. Dorsal part of the callus with straight hairs 0.2–0.5 mm long, callus 2–2.4 mm long . . .......................
....................................... S. caucasica Schmalh. var. fanica M. Nobis, P. D. Gudkova & A. Nowak
44. Longest hairs in the lower (1/3–1/2) part of lower segment of awn (column) 0.2–0.7(–0.9) mm long . . . . . . . . .....
.................................................. S. caucasica subsp. nikolai M. Nobis, A Nobis & A. Nowak
449. Longest hairs in the lower (1/3–1/2) part of lower segment of awn (column) (0.9–)1–2(–2.5) mm long . . . . . . . . . . .
............................................................................ S. caucasica subsp. caucasica
45(14). Awn pilose throughout with hairs (0.1–)0.2–7 mm long or scabrous on the first segment of the column and piloseon
the second with hairs increasing toward geniculations . . ................................................46
459. Awn glabrous or scabrous on both segments of the column and plumose on seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72
46. Upper segment of awn (seta) with hairs 0.2–1.5(–1.7) mm long . . ...........................................47
469. Upper segment of awn (seta) with hairs (1.5–)2–6.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
47. The longest ligules of vegetative shoots up to 0.3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
479. The longest ligules of vegetative shoots 0.5–2(–3) mm long . . . . .............................................55
48. Callus 0.6–1.1 mm long, awn (3.5–)4–7(–8) cm long, glumes 11–18 mm long . . . ..............................49
489. Callus 1.3–2 mm long, awn (8.5–)9–20 cm long, glumes (13–)20–40 mm long . . . . . . . . . . . . . . . . .................50
49. Seta covered with hairs 0.2–0.3(–0.5) mm long, adaxial surface of vegetative leaves with hairs 0.15–0.35 mm long; high
mountain plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. richteriana Kar. & Kir. subsp. jagnobica (Ovcz. & Czukav.) Tzvelev
499. Seta covered with hairs (0.3–)0.4–0.8(–1) mm long, adaxial surface of vegetative leaves with hairs up to 0.1 mm long, if hairs
longer than abaxial surface then densely pubescent; plants of lowlands and lower mountain belt . . . S. richteriana subsp. richteriana
50. Awn with hairs on seta 0.3–0.8(–0.9) mm long . . .........................................................51
509. Awn with hairs on seta 1–1.8 mm long . . . . . . . . .........................................................54
51. Callus 1.3–1.8 mm long, glumes 13–18 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. korshinskyi Roshev.
519. Callus more than 1.8 mm long, glumes more than 22 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
52. Awn 7–16 cm long, unigeniculate or indistinctly bigeniculate ...............................................53
529. Awn 15–20 cm long, distinctly bigeniculate . . .....................................S. 3pseudocapillata Roshev.
53. Awn 11–15 cm long, glumes 32–40 mm, vegetative leaves highly scabrous due to densely distributed spinules and
prickles ..............................................................................S. 3balkhanabatica
539. Awn 7–11 cm long, glumes 24–30 mm long, vegetative leaves glabrous or slightly scabrous due to scattered
spinules . . . . . .............................................................S. 3consanguinea Trin. & Rupr.
54(50). Anthecium 7–9.2 mm long, pilose all over with well-developed ring of hairs at the apex, awn with hairs on seta
1–1.5 mm long, callus 1.3–1.6 mm long, glumes 15–20 mm long . . . . . . . . . . . . . . . . . . . . S. 3heptapotamica Golosk.
549. Anthecium 9.5–11.5 mm long, with hairs arranged in 7 lines and without ring of hairs at the apex, awn with hairs on seta
1.4–1.8 mm long, callus 1.5–2.2 mm long, glumes 18–28 mm long . . ....................S. 3dzungarica M. Nobis
55(47). Awn with hairs on seta distinctly longer than those on column . . . ........................................56
559. Awn with hairs on seta shorter than or almost equal to those on column . . . . . ................S. roborowskyi Roshev.
56. Column with hairs up to 0.3 mm long, seta with hairs 0.4–1.1(–1.3) mm long.....................................
....................................................... S. sczerbakovii Kotukhov s.l. (including S. kotuchovii M. Nobis)
569. Column with hairs (0.1–)0.3–1 mm long, seta with hairs 1–1.5 mm long . . . . .................................57
57. Anthecium 0.5–0.7 mm wide, with 7 distinct lines of hairs with dorsal ones terminating at mid-lemma length, lemma at
the distance of ca. 1 mm below the top with a wreath of long prickles surpassed by well-developed ring of hairs at the
apex, callus densely pilose with hairs 0.3–0.5 mm long, base of callus 0.2–0.3 mm long . . . . . . . . . S. breviflora Griseb.
579. Anthecium 0.7–0.9 mm wide, irregularly pilose or with 7 more or less distinct lines of hairs with dorsal ones terminating
in 3/4–4/5 of the lemma length, lemma without a wreath of long prickles but with well-developed ring of hairs at the
apex, callus densely pilose with hairs 0.6–1 mm long, base of callus 0.4–0.7 mm long . . . . . S. 3czerepanovii Kotukhov
58(46). Panicle lax, branches flexuous, glumes pinkish violet to dark violet with hyaline margins, short (up to 5 mm),
awnlike, and sparsely ciliate cusp at the apex . . . . . . . . . ..................................S. purpurea Griseb.
8 Annals of the
Missouri Botanical Garden

589. Panicle narrow, branches erect, glumes pale green or rarely slightly purplish along the middle vein, with hyaline
margins andapex....................................................................................59
59. Longest ligules of vegetative shoots 0.2–1(–1.3) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . 60
599. Longest ligules of vegetative shoots 1.5–9(–14) mm long . . . . ...............................................61
60. Abaxial surface of leaves glabrous, anthecium 11.5–13 mm long, awn with the lower segment of column scabrous or
covered with spinules (0.05–0.2 mm long) and upper segment of the column pilose with hairs up to 2 mm long,
increasing toward the seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. 3gnezdilloi
609. Abaxial surface of leaves distinctly scabrous, anthecium 8.5–10 mm long, awn with lower segment of column pilose with
hairs up to 0.4–0.5 mm long on the lower segment and 1–1.5 mm long on the upper segment, increasing toward the
seta . . . . .........................................................................S. 3kamelinii Kotukhov
61. Anthecium 5–9(–9.5) mm long, awn 4–7.5(–9) cm long . . . . . . . . . . . . . . . ....................................62
619. Anthecium (8–)9–23 mm long, awn (9–)12–35 cm long . . . . . . . . . . . . . . . ....................................63
62. Anthecium (5–)6–7(–8) mm long, callus 0.8–1.2 mm long, top of lemma glabrous or with poorly developed ring of hairs,
awn delicate, 0.2(–0.3) mm wide near the base, with hairs on column 0.8–2.2 mm long, vegetative leaves 0.2–0.3 mm
wide and at the adaxial surface covered with prickles up to 0.1 mm long, ligules acute, ciliate only at the apex . . .
................................................................................ S. zeravshanica M. Nobis
629. Anthecium (6–)7–9(–9.5) mm long, callus (1.1–)1.3–1.6(–2) mm long, top of the lemma with well-developed ring of
hairs, awn more robust, 0.3–0.4 mm wide near the base, with hairs on the column 0.2–0.8 mm long, vegetative leaves
0.3–0.7 mm wide, and adaxial surface covered with hairs 0.15–0.25 mm long, ligules acute or obtuse, long-ciliate along
the margins .............................................................................S. orientalis Trin.
63. Upper segment of awn (seta) with hairs 1.5–2.8(–3.2) mm long . . . ..........................................64
639. Upper segment of awn (seta) with hairs (4–)4.5–6.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
64. Callus glabrous, rarely with a few scattered hairs ventrally . . . . . . . ......................S. badachschanica Roshev.
649. Callus densely pilose all over or densely pilose ventrally and with scattered hairs dorsally . . . . . ..................65
65. Awn 10–17 cm long, both segments of column pilose with hairs (0.7–)1–2 mm long, increasing
toward seta........................................................................ S. arabica Trin. & Rupr.
659. Awn 10–18 cm long, lower part of column scabrous or scabrous to shortly pilose toward first geniculation, upper
segment of column with 0.3–2 mm long hairs increasing toward second geniculation ............................66
66. Lower part of column scabrous to shortly hairy toward first geniculation, upper segment of column with 1–2 mm long
hairs increasing toward second geniculation . . .......................................S. 3assyriaca Hand.-Mazz.
669. Lower part of column scabrous, upper segment of column scabrous or with 0.3–1.5 mm long hairs increasing toward
second geniculation .........................................................S. hohenackeriana Trin. & Rupr.
67. Glumes up to 40 mm long, anthecium (8–)9–12 mm long, lemma with more or less developed ring of hairs at the apex . . .
................................................................................................... 68
679. Glumes more than 45 mm long, anthecium 12–22 mm long, lemma glabrous (without a ring of hairs) at the apex . . . 69
68. Lemma with well-developed ring of 1–3.5 mm long hairs at the apex, awn plumose throughout, column with hairs
1.5–2.5 mm long, seta with hairs 4–5 mm long, leaves of vegetative shoots 0.2–0.4 mm in diam. . . . S. himalaica Roshev.
689. Lemma with poorly developed ring of 0.05–0.4 mm long prickles at the apex, awn with lower segment of column
scabrous with mixture of 0.1–0.3 mm long prickles, upper segment pilose with hairs 0.2–0.7 mm long increasing toward
seta, seta with hairs 4.5–5 mm long, leaves of vegetative shoots (0.4–)0.5–0.7 mm in diam. . . . S. 3tzveleviana Kotukhov
69. Leaves scabrous due to prickles and spinules, lemma with dorsal and subdorsal lines of hairs fused and
terminating in lower 1/3 of the lemma length whereas ventral line terminates at 0.5–1 mm below lemma
apex.................................................S. zalesskii Wilensky var. iljinii (Roshev.) Tzvelev
699. Leaves scabrous due to prickles and hooks, lemma with dorsal and subdorsal lines of hairs not fused and terminating in
1/2–3/4 of the lemma length whereas ventral line terminates at 1.5–5 mm below the lemma apex . . . . . . . . . . . . . . . . . 70
70. Anthecium 12–14 mm long, ligules of the vegetative shoots (2–)4–6(–8) mm long.......................
............................................ S. macroglossa P. A. Smirn. var. kungeica (Golosk.) M. Nobis
709. Anthecium 14–20 mm long, ligules of the vegetative shoots (0.3–)1–2(–3) mm long . . .........................71
71. Anthecium (16–)17–19(–20) mm long, ventral line of lemma hairs terminating at 4–6(–7) mm below the apex, callus
3.5–4.5 mm long . . . . . ...................S. borysthenica Klokov ex Prokudin var. anomala (P. A. Smirn.) M. Nobis
719. Anthecium 14–16(–17) mm long, ventral line of lemma hairs terminating at 1.5–3(–4.5) mm below the apex, callus
2.5–3.5 mm long ........................S. kirghisorum P. A. Smirn. var. balkhashensis M. Nobis & P. D. Gudkova
72(45). Glumes 13–20 mm long, anthecium 5–7 mm long,awn 4.5–9 cm long, column scabrous, seta plumose . . . S. gracilis Roshev.
729. Glumes more than 21 mm long, anthecium 8.5–26 mm long, awn 10–45 cm long, column glabrous or scabrous, seta
plumose..........................................................................................73
73. Upper segment of awn (seta) with hairs 1.6–3.3 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74
739. Upper segment of awn (seta) with hairs 4–7 mm long . . ...................................................76
74. Ligules of the vegetative shoots 0.1–0.2(–0.3) mm long ............................... S. lessingiana Trin. & Rupr.
749. Ligules of the vegetative shoots (0.5–)1–6(–8) mm long . . . . . ................................................75
75. Anthecium 14–15.5 mm long, awn 19–23 cm long, column glabrous and smooth, seta with hairs 2–2.5 mmlong...
...................................................................................... S. kempirica Kotukhov
759. Anthecium 9–13(–14) mm long, awn (9–)10–15(–18) cm long, column scabrous on both segments, seta with hairs
1.6–2(–2.5) mmlong....................................................................S. hohenackeriana
76(73). Leaves of vegetative shoots setaceous, (0.1–)0.2–0.3(–0.4) mm in diam., tapering into a long bristle-like
apex ...................................................................................S. tirsa Steven
769. Leaves of vegetative shoots usually wider, at the apex with a tassel of hairs or glabrous and obtuse (but not bristle-
like) . ...............................................................................................77
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77. Lemma with dorsal and subdorsal lines of hairs fused and terminating in lower 1/5–1/3 of the lemma length, dorsal line
shorter, equal to, or slightly longer than subdorsal lines . . . . . . . . ............................................78
779. Lemma with dorsal and subdorsal lines of hairs not fused or fused only at the base and terminating in (1/3–)1/2–3/4 of
the lemma length, dorsal line 1.5–2 times longer than subdorsal lines ........................................82
78. Ventral line of hairs reaching the top of the lemma, abaxial surface of leaves scabrous due to spinules and prickles
whereas adaxial surface is covered with very short prickles (present on upper surface of ribs) and short hairs 0.2–0.3 mm
long, present only on the lateral sides of ribs .............................................................79
789. Ventral line of hairs reaching or not reaching the top of the lemma that then terminates at 0.3–5 mm below the lemma
apex, abaxial surface of leaves scabrous due to spinules and prickles or densely pilose, whereas adaxial surface is
covered with mixture of short (up to 0.15 mm long) and long (up to 0.4 mm long) hairs, present on the upper sides of
ribs . . .............................................................................................80
79. Sheaths of upper cauline leaves glabrous to slightly scabrous, ligules glabrous on the dorsal surface and ciliate on
margins, abaxial surface of leaves scabrous due to hooks and prickles . . . . . S. pulcherrima K. Koch subsp. pulcherrima
799. Sheaths of upper cauline leaves densely short pilose, ligules densely pilose on the dorsal surface and margins, abaxial
surface of leaves scabrous due to spinules and prickles . . . . . . S. pulcherrima subsp. crassiculmis (P. A. Smirn.) Tzvelev
80. Ventral line of hairs terminates at 3–5 mm below the top of the lemma . . . . . . . ............S. ucrainica P. A. Smirn.
809. Ventral line of hairs reaching the top of lemma or terminating at 0.3–1 mm below the top . . .....................81
81. Leaves of the vegetative shoots 0.5–1.2 mm in diam., abaxial and adaxial surfaces densely pilose, hairs on adaxial
surface clearly protruding from the needle-like, convoluted leaf blades . . .............S. dasyphylla (Lindem.) Trautv.
819. Leaves of the vegetative shoots 0.3–0.8 mm in diam., abaxial surface scabrous due to mixture of prickles, spinules, and
hairs, adaxial surface covered with mixture of short and long hairs, those not protruding from the needle-like, convoluted
leaf blades ...................................................................................S. zalesskii
82(77). Lemma with poorly developed ring of 0.05–0.4 mm long prickles or short hairs at the apex, awn with lower
segment of column scabrous with mixture of 0.1–0.3 mm long prickles, upper segment of column scabrous to
shortly pilose with hairs 0.2–0.7mmlongincreasingtowardthe seta.....................S. 3tzveleviana
829. Lemma without a ring of hairs at the apex, awn with lower segment of column glabrous or scabrous but without mixture of
prickles and/or hairs .................................................................................. 83
83. The longest ligules of the vegetative shoots up to 3(–3.7) mm long . . . . . ......................................84
839. The longest ligules of the vegetative shoots more than 4 mm long . . .........................................87
84. Anthecium (16–)17–19(–20) mm long, callus 3.5–4.5(–5.0) mm long, callus base narrowly cuneate . . . . . S. borysthenica
849. Anthecium 10–17(–19.5) mm long, callus 1.5–3.3(–3.8) mm long, callus base more or less pyriform . . . . . . . . . . . . . 85
85. Leaves of vegetative shoots slightly scabrous to glabrous, anthecium (16–)17–19(–19.5) mm long, ventral line of hairs
on the anthecium terminating at 4–5.5(–7) mm below the top, awn 25–40 cm long . . . . . . . . . . . . . . . . . . . . S. pennata L.
859. Leaves of vegetative shoots distinctly scabrous due to short prickles and/or spinules, anthecium (10.6–)12–16(–17) mm
long, ventral line of hairs on the anthecium terminating at (0.5–)1.2–3.1(–4.6) mm below the top, awn 14–35 cm long . . . 86
86. Anthecium (13–)14–16(–17) mm long, awn (18.5–)23–34.2 cm long, leaves on vegetative shoots 0.3–0.6 mm in diam.,
abaxial surface distinctly scabrous due to hooks and short prickles . . . . . ................S. kirghisorum P. A. Smirn.
869. Anthecium (10.6–)12–13.1(–13.8) mm long, awn (14.5–)16.3–19.6(–22.5) cm long, leaves on vegetative
shoots 0.2–0.4 mm in diam., abaxial surface distinctly scabrous due to long prickles and usually also short
hairs.......................................................................S. trichoides P. A. Smirn.
87(83). Seta 14–29 cm long, 3–9 times longer than column, glumes 42–76 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88
879. Seta 6.5–13.5 cm long, 2–4 times longer than column, glumes 25–50 mm long . . . . . . . . . . . . . . . S. turkestanica Hack.
88. Hairs on the adaxial surface of the vegetative leaves 0.15–0.25 mm long, anthecium (12.2–)13–14.5(–16) mm long,
dorsal line of hairs extending to 1/2 lemma length and terminating at (4.3–)5.5–6.8(–9.1) mm below
the top.................................................................S. macroglossa subsp. macroglossa
889. Hairs on the adaxial surface of the vegetative leaves up to 0.1 mm long, anthecium (11.3–)12–13.5(–13.9) mm
long, dorsal line of hairs extending to 2/3 lemma length and terminating at (3.1–)3.5–4.5(–6.4) mm
below the top . . . . . . . . . . . . . . . . . . . . . . . . . . . . ...............................S. macroglossa subsp. kazachstanica
Stipa 3adamii M. Nobis, Nordic J. Bot. 28(6): 734.
2010, pro sp. (S. caucasica Schmalh. 3S. kar-
ataviensis Roshev.). TYPE: Kazakhstan. Western
Tian-Shan, Karatau Mtns., upper part of the Kara-
sai Valley, Kara-sai plateau (NE of Algabas vil-
lage near Chayan), 1105 m, 12 June 1936, A.
Pyataeva s.n. (holotype, LE!; isotypes, LE!, KRA
382216!).
Habitat. Stipa 3adamii is found in mountain
steppes at 800–1200 m.
Distribution. Stipa 3adamii is found in Middle Asia
(Nobis, 2010, 2013) in Kazakhstan (Karatau Mountains;
Fig. 3).
Notes. This species was previously known only from
the type specimens; however, during revision of herbarium
materials in MW and TK, we found additional specimens
representing this nothospecies. They were collected in the
northwestern part of the Karatau Mountains, near Ran
settlement, on 13 May 1935 (G. Tekutev 85,MW),andthe
Karatau Mountains, in May 2014 (A. Ebel s.n.,TK).
Stipa akseirica Kotukhov, Turczaninowia 1(2): 11. 1998.
TYPE: Saur-Tarbagataj, praemontia boreali-occidentalia
jugi Saikan, locus Akseir, denudationes argillarum
tertiarum, partitiones glareoso-argillosae, 20 July 1993,
Ju. Kotuchov s.n. (lectotype, designated by Nobis
& Gudkova [2016: 32], LE!; isolectotypes, KRA
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432648!, KRA 436049!, KUZ!, LE!). 5Stipa
sareptana A. K. Becker.
Stipa aktauensis Roshev., Bull. Jard. Bot. Acad. Sc.
URSS 1931, 30: 302. 1932. TYPE: Turkestan.
Petro-Alexandrovsk, the Kyzyl Kum desert, Aktau,
6 May 1916, S. Filatov 219 (holotype, LE!).
Synonyms. 5Stipa lingua A. Junge var. minor
Roshev., in B. Fedtsch. Fl. Aziat. Ross. 12: 146. 1916.
TYPE: betw. Karak Ata & Adam Kirulgan, 1873, Korolkov
&Krauses.n.(holotype, LE!).
Habitat. Stipa aktauensis is found in rocky grass-
lands at 300–600 m.
Distribution. Stipa aktauensis is found in Middle
Asia (Gonzalo et al., 2012; Nobis, 2012), specifically in
Uzbekistan (Aktau Mountains). Figure 4.
Stipa 3alaica Pazij, Opred. Rast. Sred. Azii 1: 200.
1968 (S. caucasica Schmalh. 3S. trichoides P. A.
Smirn.; Nobis, 2010). TYPE: Pamiro-Alai, Alaiskii
khrebet, yugo-zap. sklon pravogo berega r. Kyzyl-su
v 50 km k zap. ot Sary-tash, na melkozemisto-
shchebenchatykh uchastkakh sredi krasnopescha-
nikovykh porod, 1 Aug. 1962, E. Puchkova 152
(holotype, TASH!).
Synonyms. 5Stipa kopetdaghensis Czopanov.
Habitat. Stipa 3alaica is found in mountain steppes
at 1500–3000 m.
Distribution. Stipa 3alaica is found in Middle Asia
(Pazij, 1968; Gonzalo et al., 2012; Nobis, 2013; Nobis
et al., 2015a), specifically in Afghanistan, Kyrgyzstan,
Tajikistan, and Turkmenistan.
Stipa 3albasiensis L.Q.Zhao&K.Guo,Ann.Bot.Fenn.
48(6): 522. 2011, pro sp. (S. glareosa P. A. Smirn. 3
S. orientalis Trin.). TYPE: China, Inner Mongolia,
Ordos, Mt. Albasi, 39°38ʹ12.10N / 106°58ʹ53.80E,
on stony mtn. slopes, alt. 1500 m, 12 June 2009,
L. Q. Zhao 09-001 (holotype, HIMC not seen).
Habitat. Stipa 3albasiensis is found in sandy
steppes, rocky grasslands, and high mountain deserts
at 2500–4000 m.
Table 1. Morphological characters and character states used in cluster analysis.
Morphological characters States
Macromorphological characters
Mean length of anthecium (lemma 1callus) (mm) 3–8.5 (1), 9–14.5 (2), 15–24 (3)
Lemma lobes absent (1), always present and rounded, triangular to elongated (2),
present and elongated with apical awnlike vein (3)
Lemma apex with hard, deflexed (retrorse) prickles (1), without hard, deflexed
prickles (2)
Mean length of callus (mm) 0.3–0.9 (1); 1–2.6 (2), 2.7–5 (3)
No. of awn geniculations without geniculation or indistinctly unigeniculate (1), unigeniculate
(2), bigeniculate (3)
Mean length of awn (mm) 4–9 (0), 10–40 (1), 41–140 (2), 141–380 (3)
Mean length of seta (mm) 3–8 (0), 9–40 (1), 41–130 (2), 131–300 (3)
Mean length of hairs on column (mm) glabrous (0), scabrous 0–0.3 (1), pubescent 0.4–2 (3), pilose 2–4 (4)
Mean length of hairs on seta (mm) 0.1–0.3 (1), 0.4–0.9 (2), 1–3 (3), 3–9 (4)
Mean length of glumes (mm) 6–17 (1), 18–38 (2), 39–85 (3)
Ratio lower glume/upper glume subequal (1), lower distinctly longer than upper (2), lower distinctly
shorter than upper (3)
Mean length of ligules of the vegetative leaves (mm) 0–0.2 (1), 0.2–1.2 (2), 1.3–3 (3), 3–8 (4)
No. of ovary styles 3 to 4 (1), 2 (2)
Micromorphological characters of the lemma
epidermis
Length of long cells 1–3(–5) times as long as width (1), (4–)5–7(–9) times as long as width
(2), wider than longer (3)
Side walls of long cells not thickened (1), thickened (2)
Hooks distribution frequent (more than 12 on area of 0.015 mm
2
) (1), sparse (less than 12
on area of 0.015 mm
2
) (2), absent (3)
Silica cells with constrictions (1), without constrictions (2)
Silica cell shape elongated to ovate (1), ovate to elliptic or reniform (2)
Silica cell distribution frequent (more than 20 per area of 0.015 mm
2
) (1), sparse (less than 20
per area of 0.015 mm
2
) (2)
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Nobis et al. 11
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Distribution. Stipa 3albasiensis is found within the
range of the parental species, S. orientalis and S.
glareosa (Mongolia, Kazakhstan, northern and western
China; Zhao & Guo, 2011; Nobis, pers. obs.). In Middle
Asia, it specifically occurs in China, Kazakhstan,
Kyrgyzstan (Tian-Shan Mountains), and Tajikistan
(Pamir).
Notes. This taxon was described from China (Zhao
& Guo, 2011). Due to intermediate characters (ligules of
vegetative shoots 0.5–1 mm, glumes 18–26 mm long,
unigeniculate or indistinctly bigeniculate awns) and the
fact that both Stipa orientalis and S. glareosa commonly
occur together in northern China, we consider
S. 3albasiensis to be a result of their hybridization.
Until this study, S. 3albasiensis was known only from
the type; however, during field studies and review of the
herbarium materials at LE, FRU, TAD, KUZ, and KRA,
we found additional specimens from Mongolia, Kazakh-
stan, Kyrgyzstan, and Tajikistan.
New records of Stipa 3albasiensis.KAZAKHSTAN. East
Kazakhstan Distr., Markakolskii region, Karaniryuk Mtns., SE
slope, 11 June 1998, O. M. Maslova et al. s.n. (KUZ!).
MONGOLIA. Gobi, Argalant Mtns., ca. 4 km of Ubchu set-
tlement, alt. 1700 m, 2 Aug. 1973, N. S. Golubkova & U. Tsogt
152 (LE!). TAJIKISTAN. Pamir Mtns., stony steppes, 10 km
SW of Murgab (hills near the rd.), alt. 3655 m, slope 25°,
inclination SE, 24 July 2016, M. Nobis & A. Nobis s.n.
(KRA!).
Stipa aliciae Kanitz, N¨ov´enyt. Gyujtesek Eredm. Grof
Szechenyi Bela Keletazsiai Utjabol 61. 1891.
TYPE: [China.] Prov. Kan-su, Ku-lang-shien, 24
June 1879, L´oczy 62 (holotype, PB not seen). 5
Stipa breviflora Griseb. (syn. after Freitag,
1985).
Stipa anomala P. A. Smirn., Del. Sem. Hort. Bot. Univ.
Mosquensis 15. 1930. TYPE: Uralskii okrug,
Teplovskii raion, mezhdu khutorami Faduleevym
i Novenkim, v zapadine sredi kovyl’noi stepi, 16
June 1929, N. Rubtsov s.n. (holotype, LE!; isotype,
MW!). [Stipa borysthenica Klokov ex Prokudin
var. anomala (P. A. Smirn.) M. Nobis.
Stipa arabica Trin. & Rupr., Sp. Gram. Stipac. 77.
1842. TYPE [original label]: 107. Stipa barbata
Desf., Unio itiner., inter lapides ad radices montis
Sinai, 15 May 1835, Schimper s.n. (lectotype,
designated here, LE TRIN, N. 1378.1!; isolecto-
types, BM!, E!, G not seen, L not seen, LE! [2
sheets], NY not seen, W!).
Notes. Stipa arabica was described by Trinius and
Ruprecht (1842) from Sinai (Egypt). The original de-
scription stated: “Inter lapides ad radices montis Sinai;
15 Majo. (Schimper! Un. itin. n. 107.).”The syntypes of
this taxon, examined by us, are preserved at LE, BM, E,
and W. The first typification of this species was
Figure 3. Distribution maps. Stipa 3adamii M. Nobis (4); S. 3fallax M. Nobis & A. Nowak (s); S. 3gnezdilloi Pazij (■);
S. gracilis Roshev. (□); S. kirghisorum P. A. Smirn. var. balkhashensis M. Nobis & P. D. Gudkova (▼); S. lingua A. Junge (:);
S. macroglossa P. A. Smirn. subsp. kazachstanica (Kotukhov) M. Nobis (=); S. magnifica A. Junge (C); S. turkestanica Hack. (à); S.
zeravshanica M. Nobis (¤).
12 Annals of the
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provided by Tzvelev (1976: 583), who stated that the
holotype and two isotypes are preserved at LE, but he
did not point out which of the specimens/sheets is the
holotype nor did he designate a lectotype. Consequently,
we designated the specimen with label “Herbarium
Trinii, N. 1378.1”as the lectotype in accordance with
the International Code of Nomenclature (McNeill et al.,
2012; Turland et al., 2018).
Within the species two varieties are recognized.
1. Leaves of vegetative shoots scabrous, rarely almost
glabrous . . . . . Stipa arabica Trin. & Rupr. var. arabica
19. Leaves of vegetative shoots pubescent . . .......
........ Stipa arabica var. turgaica (Roshev.) Tzvelev
Stipa arabica var. arabica.
Synonyms. [Stipa barbata Desf. var. arabica
(Trin. & Rupr.) Boiss. ex Kuntze, Acta Horti Petrop.
10: 255. 1887; [S. barbata var. arabica (Trin. &
Ru pr. ) Bornm., Bull. Herb. Boissier, s´er. 2, 8: 737. 1908; 5
S. caspia K. Koch [S. arabica subsp. caspia (K. Koch)
Tzvelev, Novosti Sist. Vissh. Rast. 11: 16. 1974; 5S.
szovitsiana Trin.; [S. arabica var. szovitsiana Trin. &
Rupr., Sp. Gram. Stipac. 77. 1842; [S. arabica var.
szovitsiana Trin., Mem. Acad. Petersb. ser 6, 7: 2.
1843; [S. szovitsiana (Trin. & Rupr.) Griseb., Fl.
Ross. 4: 450. 1852; 5S. orientalis Trin. var. persica
Trin., M ´em. Acad. Imp. Sci. St. P ´etersbourg Hist. Acad.
1: 79. 1830 (TYPE: LE? not seen).
Habitat. Stipa arabica var. arabica isfoundinstony
and sandy steppes, fallows, and roadsides at 400–2800 m.
Distribution. Stipa arabica var. arabica is widely
distributed from Asia Minor to central Asia (Tzvelev,
1976, 2006; Freitag, 1985; Wu & Phillips, 2006; Nobis
et al., 2016b). In Middle Asia, it is found in Afghani-
stan, China, Iran, Kazakhstan, Kyrgyzstan, Pakistan,
Tajikistan, Turkmenistan, and Uzbekistan.
Stipa arabica var. turgaica (Roshev.) Tzvelev, Zlaki
SSSR 584. 1976. Basionym: Stipa turgaica
Roshev., Bot. Mater. Gerb. Bot. Inst. Komarova
Akad. Nauk S.S.S.R. 11: 21. 1949.
Synonyms. 5Stipa damascena Boiss.; 5S. koeni-
gii Woronow; [S. arabica Trin. & Rupr. subsp.
koenigii (Woronow) Tzvelev, Bot. Zhurn. (Moscow &
Leningrad) 78(10): 93. 1993.
Habitat and distribution. The range and habitats for
Stipa arabica var. turgaica are similar to those of
S. arabica var. arabica.
Notes. Stipa arabica s.l. is highly variable in mor-
phology, and a taxonomic revision of its specimens from
the entire distribution range is needed.
Stipa 3assyriaca Hand.-Mazz., Ann. Nat. Hofmus.
Wien 28: 26. 1914, pro sp. (S. arabica Trin. & Rupr.
3S. hohenackeriana Trin. & Rupr.). TYPE: [Iraq.]
Gipssteppe auf dem R¨ucken des Kalaat Schergat
Figure 4. Distribution maps. Stipa aktauensis Roshev. (4); S. 3czerepanovii Kotukhov (▼); S. 3heptapotamica Golosk. (=);
S. himalaica Roshev. (à); S. karataviensis Roshev. (s); S. longiplumosa Roshev. (□); S. narynica M. Nobis (C); S. okmirii Dengub.
(■); S. richteriana Kar. & Kir. subsp. jagnobica (Ovcz. & Czukav.) Tzvelev (
¤); and S. tianschanica Roshev. (:).
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Nobis et al. 13
Synopsis of Stipa (Poaceae) in Middle Asia

(Assur) am Tigris, 200–300m,10May1910,
Handel-Mazzetii 1062 (holotype, WU 0072644!).
Synonyms. [Stipa hohenackeriana Trin. & Rupr.
var. assyriaca (Hand.-Mazz.) H. Scholz, Fl. Turkey 9:
548. 1985; [S. hohenackeriana subsp. assyriaca
(Hand.-Mazz.)F.M.V´azquez, Telopea 13: 168. 2011; 5
S. stapfii Roshev.; 5S. iraqensis Martinovsk´y; 5
S. 3kolakovskyi Tzvelev.
Habitat. Stipa 3assyriaca is found in sandy steppe,
rocky grasslands, and roadsides at 400–1500 m.
Distribution. Stipa 3assyriaca is found within the
range of both parental species, S. arabica and S.
hohenackeriana. In Middle Asia, it occurs in Afghani-
stan, northern Iran, Kazakhstan, Kyrgyzstan, Tajikistan,
Turkmenistan, and Uzbekistan.
Stipa atriseta Stapf ex Bor, Fl. Iranica [Rechinger], 70:
389. 1970. TYPE: [Iran, Fars Prov.] In summo
monte Kuh Bul, 4600 m, 6 Sep. 1885, Stapf 1536
(holotype, W!). 5Stipa hohenackeriana Trin. &
Rupr. var. hohenackeriana.
Stipa avenoides Honda, Rep. Exped. Manchoukuo Sect.
IV, Pt. 4, Index Fl. Jeholensis 103. 1936. TYPE: not
seen. 5Achnatherum sibiricum (L.) Keng ex
Tzvelev, Probl., Ekol. Geobot., Bot. Geogr. Florist.
140. 1977 (syn. after Wu & Phillips, 2006).
Stipa azutavica Kotukhov,Turczaninowia 1(2): 9. 1998.
TYPE: Altaj australis, praemontia australi-orientalia
jugi Azutau, mons Bulgartabaty, desertum lapido-
sum, 22 May 1991, Ju. Kotuchov s.n. (lectotype,
designated by Nobis & Gudkova [2016: 36], LE!;
isolectotypes, KRA 436050!, KUZ!, LE!). [Stipa
orientalis Trin. var. azutavica (Kotukhov) M. Nobis
&P.D.Gudkova.
Stipa badachschanica Roshev., Bot. Mater. Gerb.
Bot. Inst. Komarova Akad. Nauk S.S.S.R. 11: 19.
1949. TYPE: [USSR, Pamir] Schugnan, dolina
Darshaz ot mosta Pul-i-furs do ust’ya, 9 June
1914, N. Tuturin & P. Bessedin 183 (lectotype,
designated by Tzvelev [1976: 584], LE!; isolec-
totypes, LE!).
Within the species two varieties are recognized.
1. Lemma covered with scattered hairs . . . . . . . . . . .
... Stipa badachschanica Roshev. var. badachschanica
19. Lemma glabrous, rarely with single hairs in the
lower part . . . ..............................
Stipa badachschanica var. pamirica (Roshev.) M. Nobis
Stipa badachschanica var. badachschanica.
Habitat. Stipa badachschanica var. badachschan-
ica is found in sandy, rocky, and steppe grasslands at
1800–4000 m.
Distribution. Stipa badachschanica var. badach-
schanica is found in southwestern Asia (Ovchinnikov
& Chukavina, 1957; Tzvelev, 1976; Ikonnikov, 1979;
Freitag, 1985). In Middle Asia, it occurs in Afghanistan,
northern Iran, and Tajikistan (Pamir).
Notes. Stipa badachschanica var. badachschanica
is morphologically similar to S. arabica var. turgaica,
but can be distinguished by having a glabrous (versus
pilose) callus.
Stipa badachschanica var. pamirica (Roshev.) M.
Nobis, comb. & stat. nov. Basionym: Stipa pamir-
ica Roshev., Bot. Mater. Ger. Bot. Inst. Komarova
Akad. Nauk S.S.S.R. 11: 20. 1949. TYPE: [USSR,
Tajikistan.] Vakhan-Inkashimskii raion, kovyl’no-
solyankovo-polynnaya polupustynya po vost.
shchebnistym sklonam v raione kishlaka Vrang,
3120 m, 9 Aug. 1935, P. Ovchinnikov & K.
Afanasev 1735 (lectotype, designated by Tzvelev
[1976: 584], LE!).
Synonyms. [Stipa badachschanica Roshev. subsp.
pamirica (Roshev.) Tzvelev, Novosti Sist. Vyssh. Rast.
11: 16. 1974; [S. arabica Trin. & Rupr. subsp. pamirica
(Roshev.) F. M. V ´azquez, Telopea 13(1–2): 168. 2011; [
S. arabica var. pamirica (Roshev.) Freitag, Notes Roy.
Bot. Gard. Edinburgh 42(3): 461. 1985.
Habitat. Stipa badachschanica var. pamirica is
found in rocky and steppe grasslands and screes at
2500–4500 m.
Distribution. Stipa badachschanica var. pamirica is
found in Middle Asia (Ovchinnikov & Chukavina, 1957;
Tzvelev, 1976; Ikonnikov, 1979; Freitag, 1985), specif-
ically in Afghanistan and Tajikistan (Pamir).
Notes. Stipa badachschanica var. pamirica differs
from S. badachschanica var. badachschanica only in the
character of the lemma, which is glabrous or almost so
versus covered with scattered hairs, respectively. Plants
belonging to S. badachschanica var. pamirica were
collected from the same area as those belonging to
variety badachschanica. A similar phenomenon has
been seen in other species that normally have pubescent
lemmas, e.g., S. subsessiliflora or S. klemenzii Roshev.
(M. Nobis, pers. obs.).
Stipa baktashevae Tzvelev, Novosti Sist. Vyssh. Rast.
45: 7. 2014. TYPE: Republic of Kalmykia. Tse-
linnyi Distr., outskirts of the Solnechnyi settle-
ment, sandy slope among psammophytes, 21 May
2011, E. Egorova & N. Baktasheva s.n. (holotype,
14 Annals of the
Missouri Botanical Garden

LE 01009340!). 5Stipa borysthenica Klokov ex
Prokudin var. borysthenica.
Stipa 3balkanabatica M. Nobis & P. D. Gudkova,
nothosp. nov. (S. caucasica Schmalh. 3S. sarep-
tana A. K. Becker). TYPE: Turkmenskaya SSSR
[Turkmenistan]. khr. Bol’shoi Balkhan, tropa ot
rodn, Arlan k rodn, Kabak-Bulak, pologii sklon
melkozem., v bol’shom kolichestve [Great Balkhan
Mtns., rte. from Arlan to Kabak-Bulak, on slope,
abundant] 1600 m, 25 June 1958, Praskuryanova
s.n. (holotype, MW!; isotype, KRA476853!). Figure 5.
Diagnosis. Stipa 3balkanabatica M. Nobis & P. D. Gudkova
is similar to S. 3consanguinea Trin. & Rupr., a hybrid be-
tween S. glareosa P. A. Smirn. and S. krylovii Roshev., in its
unigeniculate awns that are minutely pubescent through-
out, but it differs in having more robust, longer (11–15 cm
vs. 7–11 cm) awns and glumes (32–40 vs. 24–30 mm) as
well as vegetative leaves that are distinctly scabrous due to
densely distributed spinules and prickles versus vegeta-
tive leaves glabrous or slightly scabrous due to scattered
spinules.
Plants perennial, densely tufted, with a few culms
and numerous vegetative shoots; culms 40–60 cm tall,
3-noded, glabrous at nodes and very densely and shortly
pubescent below them. Leaves of vegetative shoots:
sheaths glabrous, ciliate at margins and with white
edge; ligules rounded or truncate, up to 0.2 mm and
ciliate at margins; blades convolute, up to 20 cm,
0.5–0.7 mm in diam., upper surface densely pubescent
with hairs 0.15–0.2 mm, and on marginal ribs with
mixture of longer hairs 0.2–0.3 mm, lower surface
scabrous due to short prickles and bristles. Cauline
leaves: sheaths glabrous and with white edge, shorter
than internodes, upper sheath up to 20 cm, glabrous,
slightly inflated; ligules 2–3.3 mm, obtuse or acute;
blades scabrous, up to 10 cm. Panicle 17–24 cm
contracted, at base enclosed by sheath of uppermost
leaf, branches erect, setulose, single or paired. Glumes
subequal, 32–40 mm, narrowly lanceolate, tapering into
long hyaline apex, midvein sometimes setulose with
cilia up to 1 mm. Anthecium 10–12 30.7–0.8 mm.
Callus 1.9–2.2 mm, densely long-pilose on ventral and
dorsal surfaces, callus base not enlarged, peripheral
ring 0.2 mm in diam., acute, cuneate, scar elliptic to
circular. Lemma pale green, on dorsal surface with abun-
dant hooks and with 7 lines of ascending hairs, hairs up to
0.5–0.6 mm, ventral and dorsal line of hairs terminating at
1–2 mm below top of lemma; top of lemma scabrous due to
prickles and short hairs, surpassed by a ring of unequal
Figure 5. Stipa 3balkanabatica M. Nobis & P. D. Gudkova. —A. Vegetative shoots and panicle. —B. Top of the lemma. —C.
Lemma epidermal pattern. —D. Callus. —E. Ring (foot) of the callus. —F. Abaxial surface of leaves. —G. Adaxial surface of
leaves.
Volume 105, Number 1
2020
Nobis et al. 15
Synopsis of Stipa (Poaceae) in Middle Asia

hairs 0.3–1 mm long at apex. Palea equal to lemma in
length. Awn 130–160 mm, unigeniculate or slightly
bigeniculate; column 32–46 mm, twisted, 0.4–0.5 mm
wide at base, covered with hairs 0.2–0.5 mm, gradually
increasing in length toward geniculation; seta arcuate
or flexuous, 100–120 mm, hairs in lower part of seta
0.4–0.8(–0.9) mm, gradually decreasing in length toward
apex. Anthers yellow, ca. 7 mm, glabrous.
Phenology. Stipa 3balkanabatica flowers from May
to June.
Habitat. Stipa 3balkanabatica is found in moun-
tain steppes, slopes, and screes at 1500–1700 m.
Distribution. Stipa 3balkanabatica is found in
Turkmenistan (northwestern Kopet-Dagh).
Etymology. The name of the species originates from
Balkanabat, a city located in the vicinity of the Great
Balkhan Mountains.
Stipa basiplumosa Munro ex Hook. f., Fl. Brit. India
[J. D. Hooker] 7(22): 229. 1896. TYPE: Western
Tibet (India, Kashmir, Ladakh), ascent to Lank
pass, 13 Sep. 1847, Thomson s.n. (syntype, K);
ravine below Lank pass, 5000 m, 14 Sep. 1847,
Thomson s.n. (syntypes, E!, K!).
Synonyms. [Stipa subsessiliflora (Rupr.) Roshev.
subsp. basiplumosa (Munro ex Hook. f.) D. F. Cui, Fl.
Xinjiangensis 6: 309. 1996; [S. subsessiliflora var.
basiplumosa (Munro ex Hook. f.) P. C. Kuo & Y. H. Sun,
Fl. Reipubl. Popularis Sin. 9(3): 284. 1987.
Habitat. Stipa basiplumosa is found in high moun-
tain deserts and semideserts, steppes, and stony slopes
at 3000–4600 m.
Distribution. Stipa basiplumosa is found in the
mountains of central Asia (Tzvelev, 1968; Kuo &
Sun, 1987; Nobis et al., 2015b, 2016b). In Middle
Asia, it occurs in southwestern China and Pakistan.
Notes. In the Himalayas, Kunlun, and Karako-
rum, the taxon can be confused with Stipa subsessi-
liflora, but the two species clearly differ in length
and shape of glumes as well as in awn length (see
key).
Stipa bella Drobow, Repert. Spec. Nov. Regni Veg. 21:
37. 1925, hom. illeg. non Phil., Anales de la
Universidad de Chile 26: 203. 1870. [Stipa
drobovii (Tzvelev) Czerep. var. drobovii.
Stipa bhutanica Noltie, Edinburgh J. Bot. 56(2): 289.
1999. TYPE: Bhutan. Ha Distr., W side of Chel ai La,
27°229N, 89°209E, 3600 m, 29 Sep. 1998, Noltie,
Pradham, Sherub & Wangdi 349 (holotype, E!;
isotype, THIM not seen). [Ptilagrostis bhutanica
(Noltie) M. Nobis, PhytoKeys 128: 109. 2019.
Stipa boczantzevii Tzvelev, Novosti Sist. Vyssh. Rast.
43: 24. 2011 [2012]. TYPE: Asia Media, jugum
Alaicum, secus fl. Aksu inter pag. Jordan et
trajectum Shivali, in calcareis, 17 Aug. 1962, V. P.
Boczantzev 208 (holotype, LE!). 5Stipa caucasica
Schmalh. var. caucasica.
Stipa borysthenica Klokov ex Prokudin, Fl. Kryma
[Wulff] 1(4): 25. 1951. TYPE: Ukraine. Lugov
forest country house, Tyasmin, Aleks, u., 18 May
1911, I. Paczoski s.n. (lectotype, designated by
Tzvelev [1976: 591], LE!).
Within the species two varieties are recognized.
1. Awn glabrous on column and plumose on seta . . .
Stipa borysthenica Klokov ex Prokudin var. borysthenica
19. Awn pilose throughout . . . . . ..................
Stipa borysthenica var. anomala (P. A. Smirn.) M. Nobis
Stipa borysthenica var. borysthenica.
Synonyms. [Stipa pennata L. fo. sabulosa Pacz.,
Kherson.Fl.1:112.1914;[S. sabulosa (Pacz.) Sljuss.; [
S. pennata subsp. sabulosa (Pacz.) Tzvelev, Novosti Sist.
Vyssh. Rast. 10: 80. 1973; [S. joannis ˇ
Celak. subsp.
sabulosa (Pacz.) Lavrenko, Flora SSSR 2: 123. 1940; 5S.
baktashevae Tzvelev; 5S. joannis var. marchica Endtm.,
Wiss. Z. Ernst-Moritz-Arndt-Univ. Greifswald, Math.-
Naturwiss. Reihe 11: 148. 1976. TYPE: Germany.
Naturschutzgebiet “Geesower H¨ugel”zwischen Gartz/
Oder und Geeson/Kr. Angerm¨unde, 18 June 1960,
Endtmann (holotype, GFW; isotype, JE!); [S. borys-
thenica Klokov ex Prokudin var. marchica (Endtm.)
Rauschert, Mitt. Arbeitsgen. Florist. Kart. Bayerns 4:
11. 1978.
Habitat. Stipa borysthenica var. borysthenica is found
in sandy grasslands at 0–900 m.
Distribution. Stipa borysthenica var. borysthenica is
widely distributed from central Europe to central Asia
(Pazij, 1968; Tzvelev, 1968, 1976, 2006; Martinovsk´y,
1980; Freitag, 1985; Gonzalo et al., 2013; Nobis et al.,
2016b). In Middle Asia, it is found in northwestern
China and Kazakhstan.
Stipa borysthenica var. anomala (P.A.Smirn.)
M.Nobis,comb.&stat.nov.Basionym:Stipa
anomala P. A. Smirn., Del. Sem. Horti. Univ.
Mosquensis 15. 1930.
Synonyms. [Stipa anomala P. A. Smirn. ex
Roshev., Fl. URSS 2: 740. 1934; [Stipa pennata L.
var. anomala (P. A. Smirn) Tzvelev, Novosti Sist. Vyssh.
Rast. 11: 18. 1974; [Stipa pennata subsp. anomala
16 Annals of the
Missouri Botanical Garden

(P. A. Smirn.) F. M. V´azquez & M. Gut., Telopea
13(1–2): 159. 2011.
Habitat. Stipa borysthenica var. anomala is found
in sandy grasslands at 0–900 m.
Distribution. Stipa borysthenica var. anomala is found
within the range of the nominal variety (Roshevitz, 1934;
Tzvelev, 1976). In Middle Asia, it occurs in Kazakhstan.
Notes. Although this taxon belongs to Stipa sect.
Stipa, which comprises plants having awns with glabrous
columns, sometimes within populations of this species a
few individuals with shortly pubescent columns can be
recorded. According to Tzvelev (1976) and Scholz (1985),
such individuals with pubescent awn columns could be the
result of spontaneous mutation. Nevertheless, such indi-
viduals have been observed in subsequent years or con-
firmed at particular localities several years after their first
finding (e.g., S. zalesskii var. iljinii;S. pulcherrima var.
paradoxa A. Junge ex Roshev.) [synonyms: Stipa grafiana
Steven var. paradoxa A. Junge, Izv. Imp. S.-Peterburgsk.
Bot. Sada 10: 1, 1910; S. paradoxa (A. Junge) P. A. Smirn.,
Tabl. dla opred. kovyl. 7. 1927, nom. illeg. hom. non S.
paradoxa (L.) Raspail, Ann. Sci. Nat. (Paris) 5: 449. 1825;
S. syreistschikowii P. A. Smirn., Del. Sem. Hort. Bot. Univ.
Mosquensis 1948: 36. 1948; S. schisensis Roshev. ex
Grossh., Fl. Kavkaza 1: 65. 1928]. Here we treat such
specimens in the rank of variety, with the most similar
species having glabrous lower parts of the awn (column).
It is worth mentioning that while examining herbar-
ium materials from Armenia, we found another spe-
cimen from section Stipa with entirely pilose awns.
Having lemmas with seven lines of ascending hairs
0.8–1.2 mm long, of which one ventral lemma termi-
nates 0.5–1.5(–2) mm below the top of the lemma,
whereas the dorsal terminates at 4/5–3/4 of the lemma
length, about (1.5–)2–5(–6) mm below the top and being
twice as long as the subdorsal lines, these specimens are
most similar to S. araxensis; however, their entirely
pilose awns are also similar to those of S. pulcherrima
var. paradoxa. We consider that these specimens may
be the result of spontaneous mutation. They represent a
new variety of S. araxsensis, which is described below:
Stipa araxensis Grossh. var. mikojanovica M. Nobis, var.
nov. TYPE: Stipa araxensis Gross. 3S. mayeriana
Trin & Rupr. [det. N. Tzvelev], Armenian SSR,
Mikoyanovski Distr., rocky slope (almost scree),
western exposition of not high mtn. betw. Agi-Sofi
Mtns. & Arpi-Gai River, 2.5–3 km to the SSW from
the Mikoyan city [now Ehegnadzor], 7 July 1957, N.
N. Tzvelev & S. K. Cherepanov 745 (holotype, LE!).
Diagnosis. Stipa araxensis Grossh. var. mikojanovica M.
Nobis differs from S. araxensis var. araxensis in having the awn
column shortly pubescent (covered with 0.3–1 mm long hairs)
versus awn column glabrous and smooth, respectively. Due to
having entirely pilose awns it resembles S. pulcherrima K.
Koch var. paradoxa A. Junge ex Roshev.; however, it differs in
general pattern of lemma pilosity. In S. araxensis var. mikoja-
novica ventral lines terminate 0.5–1.5(–2) mm below the top of
the lemma and the dorsal line terminates in 4/5–3/4 of the
lemma length, whereas in S. pulcherrima var. paradoxa the
dorsal line of hairs is fused with subdorsals with the ventral
lines reaching the top of the anthecium.
Stipa brandisii Mez, Repert. Spec. Nov. Regni Veg. 17:
207. 1921. TYPE: [India] N. W. Himalaya, Kulla,
Oct. 1876, Dr. Brandis 1005 (lectotype, selected
and labeled by H. Freitag on 15 Mar. 1984 and
designated by Nobis et al. 2019b: 110, K
32092!) 5Achnatherum brandisii (Mez) Z. L.
Wu, Acta Phytotax. Sin. 34: 154. 1996.
Stipa brauneri (Pacz.) Klokov, Novosti Sist. Vyssh.
Nizsh. Rast. 1975: 21. 1976. Basionym: Stipa
lessingiana Trin. & Rupr. subsp. brauneri Pacz.,
Zap. Krymsk. Obsch. Estesvoisp. 5: 4. 1916.
TYPE: Tauria, penins, Tarchankut, prope Ak-
Meczet, 30 Apr. 1914, A. Brauner s.n. (holotype,
LE!). [Stipa lessingiana Trin. & Rupr. var.
brauneri (Pacz.) Roshev.
Stipa 3brevicallosa M. Nobis, Pl. Syst. Evol. 299(7):
1340. 2013 (S. lipskyi Roshev. 3S. drobovii
(Tzvelev) Czerep.; Nobis, 2013). TYPE: Dolina
Fan (Fan River valley), G ´ory Zerawsza ´nskie
(Pamiro-Ałaj), “Rejon Zerawsza ´nski C,”Dolina
rzeki Fan, prawy stok g´or Gushty, ok. 500 m na
E od centrum przysi´ołka Zerawszan, wysokog´orski
step ostnicowy [Zeravshan Mtns., Fan River Val-
ley, right slope of Gushty Mtns., ca. 500 m E of
Zeravshan settlement center, high mtns. feather
grass steppe], 39°139140N / 68°329510E, alt.
2150 m, exp. W, slope 25°, loc. 1, 21 June
2009, M. Nobis & A. Nowak s.n. (holotype, Herb.
Stip. M. Nobis!, KRA 383192!; isotypes, KRA
423399!, KRA 383490!, KRA 383084!).
Within the species two varieties are recognized.
1. Leaves of vegetative shoots glabrous . . . . . . . . . . .
........ Stipa 3brevicallosa M. Nobis var. brevicallosa
19. Leaves of vegetative shoots densely pubescent . . .
Stipa 3brevicallosa var. hissarensis M. Nobis & A. Nowak
Stipa 3brevicallosa var. brevicallosa.
Habitat. Stipa 3brevicallosa var. brevicallosa is
found in mountain steppes at 1300–2350 m.
Distribution. Stipa 3brevicallosa var. brevicallosa
is found in Middle Asia (Nobis, 2013), specifically in
Tajikistan (Hissar and Zeravshan Mountains).
Stipa 3brevicallosa var. hissarensis M. Nobis & A.
Nowak, Acta Mus. Siles. Sci. Nat. 65: 278. 2016.
Volume 105, Number 1
2020
Nobis et al. 17
Synopsis of Stipa (Poaceae) in Middle Asia

TYPE: Tajikistan. Hissar Mtns. (Pamir Alai), “Zer-
avshan B region,”steppe grassland on the right
slope of Gabierud River valley, near Pormin set-
tlement near Zeravshan village, ca. 2 km S of
Jagnob River, alt. 1896 m, 39°10912.460N/
68°34942.860E, 25 May 2015, M. Nobis & A.
Nowak s.n. (holotype, KRA 464720!; isotypes,
KRA 464721!, KRA 464722!, KRA 464723!,
KRA 464708!).
Habitat. Stipa 3brevicallosa var. hissarensis is
found in mountain steppes at 1850–2350 m.
Distribution. Stipa 3brevicallosa var. hissarensis is
found in Middle Asia (Nobis & Nowak, 2016), specif-
ically in Tajikistan (Hissar Mountains).
Stipa breviflora Griseb., Nachr. K¨onigl. Ges. Wiss.
Georg-Augusts-Univ. 82. 1868. TYPE: Tibet.
Gnari Khorsum Prov.: Peti via Lom´orti to Poling,
5–15 Sep. 1855, Schlagintweit 7105 (holotype,
GOET!; isotype, LE!).
Synonyms. 5Stipa aliciae Kanitz.
Habitat. Stipa breviflora is found in sandy steppes,
grasslands, and screes at 1000–4000 m.
Distribution. Stipa breviflora is a widely distributed
central Asian species (Tzvelev, 1968, 1976; Pazij,
1968; Freitag, 1985; Dickor´e, 1995; Lu & Wu, 1996;
Wu & Phillips, 2006). In Middle Asia, it is found in
China, Kyrgyzstan, and Pakistan.
Stipa bromoides (L.) D¨orfl., Herb. Norm. Sched. Cent.
[D¨orfler] 34: 129, no. 3386. 1897. Basionym:
Agrostis bromoides L., Mant. Pl. 30. 1767. TYPE:
Montpellier, Gouan 3 (lectotype, designated by
Freitag [1985: 401], LINN 94.6!). [Achnathe-
rum bromoides (L.) P. Beauv., Ess. Agrostogr.
20: [146], 147. 1812 [[Aristella bromoides (L.)
Bertol., Fl. Ital. [Bertoloni] 1: 690. 1833].
Stipa 3brozhiana M. Nobis, Nordic J. Bot. 29(4): 459.
2011 (S. lipskyi Roshev. 3S. arabica Trin. &
Rupr.; Nobis, 2011). TYPE: G ´ory Zerawsza ´nskie
(Pamiro-Alaj), “Rejon Zerawsza´nski B”Zeravszan
Mtns., “Zeravshanski range B”Vashan River Val-
ley (near Urmetan), murawa stepowa w cze
˛´sci podszc-
zytowej lewego zbocza doliny rzeki Vashan (w pobli˙zu
uj´scia do rzeki Zerawszan), przy N cze
˛´sci wsi Vashan
koło Urmetan (pomie
˛dzy rzeka
˛Vashan a przełe
˛cza
˛
Chukalik) [Tajikistan, Zeravshan Mtns. (Pamir Alai
Mtns.), steppe grassland near the top of the left slope
of the Vashan River valley, N of Vashan settlement
near Urmetan village (betw. the Vashan River &
Chukalik pass)], N 39°249470/ E 68°169230,1680m,
exp. NE, slope 30°, 21 June 2010, M. Nobis
(holotype, KRA 378169!; isotypes, Herb. Stip. M.
Nobis!, KRA 378170!, KRA 378171!, KRAM!).
Habitat. Stipa 3brozhiana is found in steppe grass-
lands at 1600–1800 m.
Distribution. Stipa 3brozhiana is found in Middle
Asia (Nobis, 2011, 2013), specifically in Tajikistan
(Zeravshan Mountains).
Stipa bungeana Trin. ex Bunge, Enum. Pl. China Bor.
[A. A. von Bunge] 70. 1833. TYPE: Ad radices
montium Zui-wey-schan et ad vias prope Ssi-jui-ssi
(lectotype, designated here, LE Herbarium Trinii
1383.1!; isolectotype, LE Herbarium Trinii 1382.2!;
syntypes, LE! [3 sheets]).
Habitat. Stipa bungeana is found in sandy steppes,
grasslands, and semideserts at 500–4000 m.
Distribution. Stipa bungeana is a widely distributed
central Asian species (Pazij, 1968; Tzvelev, 1968,
1976; Lu & Wu, 1996; Wu & Phillips, 2006). In Middle
Asia it is found in central Afghanistan, China, Kyrgyz-
stan (around Issyk-Kul Lake), and Pakistan.
Notes. In LE there are five sheets with specimens of
Stipa bungeana originating from the collection of Bunge.
However, only on two of them (preserved in the Trinius
collection) is the label “Ad radices montium Zui-wey-schan
et ad vias prope Ssi-jui-ssi”as stated in the protologue
(Bunge, 1833). The first typification of this species was
provided by Tzvelev (1976: 578), who stated that the ho-
lotype and three isotypes are preserved at LE, but he did not
point out which of the specimens/sheets is the holotype nor
did he designate a lectotype. Consequently, we designated
the specimen with label “Herbarium Trinii, N. 1383.1”as
the lectotype in accordance with the International Code of
Nomenclature (McNeill et al., 2012; Turland et al., 2018).
Stipa canescens P. A. Smirn. ex Roshev., Fl. URSS 2:
741. 1934. TYPE: Armenia. Distr. Nor-Bajazet: in
viciniis pag. Elenovka, 2 July 1929, O. Zedelmejer
& T. Gejdeman s.n. (holotype, LE!). 5Stipa
zalesskii Wilensky subsp. zalesskii.
Stipa capensis Thunb., Prodr. Pl. Cap. 1: 19. 1794.
TYPE: Promontorium Bonae Spei Africae, Thun-
berg 2560 (holotype, UPS not seen). [Stipellula
capensis (Thunb.) R¨oser & Hamasha, Schlech-
tendalia 24: 92. 2012 [[Stipella capensis (Thunb.)
R¨oser & Hamasha, Pl. Syst. Evol. 298: 365. 2012,
hom. illeg. [Stipella capensis (Thunb.) Tzvelev,
Novosti Sist. Vyssh. Rast. 43: 23. 2011 [2012],
hom. illeg.].
Stipa capillata L., Sp. Pl., ed. 2, 1: 116. 1762. TYPE:
Bohemia, Busser I: 127(1) (lectotype, designated
by Freitag [1985: 453], UPS not seen).
18 Annals of the
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Synonyms. 5Stipa ukranensis Lam.
Habitat. Stipa capillata is found in sandy steppes
and grasslands at 500–2500 m.
Distribution. Stipa capillata is widely distributed
from western Europe to central-eastern Asia (Pazij,
1968; Tzvelev, 1968, 1976; Martinovsk´y, 1980; Freitag,
1985; Wu & Phillips, 2006). In Middle Asia it is found
in Afghanistan, China, Iran, Kazakhstan, Kyrgyzstan,
Pakistan, Tajikistan, Turkmenistan, and Uzbekistan.
Notes. Kotukhov (1987) described Stipa austroal-
taica Kotukhov from the southern Altai Mountains,
which is morphologically very similar to S. capillata.
He distinguished it from S. capillata by its shorter anthecia
(7.5–8.2 vs. 10–13 mm long), shorter awns (8.1–9.6 vs.
11–20 cm long), and sheaths of culm leaves that are shorter
(vs. longer) than the internodes. After reviewing all speci-
mens of S. austroaltaica from the type collections at LE, we
found the variability in the length of these characters to be
greater than that described in the protologue. For example,
anthecia are 7.8–11 mm long and awns are 8.5–10.3 cm
long (Nobis & Gudkova, 2016). In S. capillata, sheaths are
generally longer than or equal to the internodes; however,
during our review of specimens, we also saw sheaths that
were somewhat shorter than internodes. It is likely that S.
austroaltaica is conspecific with S. capillata and represents
only marginal variability of the latter taxon. Taxonomic
revision of this group of species is required.
Stipa caragana Trin., M´em. Acad. Imp. Sci. St.-
P´etersbourg, S ´er. 6, Sci. Math. 1: 74. 1830. TYPE:
Littora orientalia maris Caspium ad prom. Tjuk-
Caragan, Eichwald s.n. (lectotype, designated by
Tzvelev [1976: 564], LE!). [Achnatherum caragana
(Trin.) Nevski, Trudy Bot. Inst. Akad. Nauk S.S.S.R.,
Ser. 1, Fl. i Sist. Vyssh. Rast. 4: 336–337. 1937.
Stipa caspia K. Koch, Linnaea 21(4): 440. 1848. TYPE:
Am Ufer des Caspischen Meeres, zwischen Baku und
Derbend, auf Meersand, K. Koch s.n. (lectotype,
designated here, GOET!; holotype, preserved at B
Figure 6. A comparison of selected morphological characters in Stipa caucasica Schmalh. agg. —A. Awns with anthecia. I, S.
caucasica subsp. nikolai M. Nobis, A. Nobis & A. Nowak; II, S. caucasica var. fanica M. Nobis, P. D. Gudkova & A. Nowak; III, S.
caucasica var. caucasica.—B. Awn column: S. caucasica subsp. nikolai (left) and S. caucasica var. caucasica.—C–E. Adaxial
surface of the vegetetive leaves in S. caucasica subsp. nikolai,S. caucasica var. fanica, and S. caucasica var. caucasica,
respectively. Scale bars: A 51 cm, B 50.1 mm, C–E50.3 mm.
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Nobis et al. 19
Synopsis of Stipa (Poaceae) in Middle Asia

was destroyed during World War II). 5Stipa arab-
ica Trin. & Rupr. var. arabica.
Stipa caucasica Schmalh., Ber. Deutsch. Bot. Ges. 10:
293. 1892. TYPE: Dagestan, Temir-Khan-Shura,
6 May 1891, V. Lipskii s.n. (lectotype, designated
by Tzvelev [1976: 593], LE!; isolectotypes, LE!;
ERE 418! electr. image).
Stipa caucasica subsp. caucasica.
Within the taxon, two varieties are recognized.
1. Abaxial surface of leaves densely covered with ca.
0.1 mm long prickles, callus at the dorsal surface
densely and long pilose, with hairs 1–1.5mmlong...
................. Stipa caucasica Schmalh. var. caucasica
19. Abaxial surface of leaves densely covered with
0.25–0.5 mm long hairs, callus at the dorsal surface
sparsely pilose, with short (0.4–0.8mm long) hairs . . .
...................................... Stipa
caucasica var. fanica M. Nobis, P. D. Gudkova & A. Nowak
Stipa caucasica var. caucasica (Fig. 6).
Synonyms. 5Stipa orientalis Trin. var. grandiflora
Rupr., M´em. Acad. Imp. Sci. Saint-P´etersbourg, S ´er. 7,
14(4): 35. 1869. TYPE: Kyrgyzstan. In regione sylvatica
jugi Tian-Shan, 20 July 1867, Osten-Sacken s.n. (lec-
totype, designated by Gonzalo et al. [2011: 401], LE!);
5S. caucasica fo. autumnalis Roshev., Fl. Aziat. Ross.
12: 142. 1916. TYPE: Kazakhstan. Prov. Semirichinsk,
Distr. Djarkinsk, Kapkak River, 13 July 1912, Sap-
oshnikow & Schischkin s.n. (lectotype, designated by
Gonzalo et al. [2011: 401], LE!); 5S. caucasica fo.
crassifolia Roshev., Fl. Aziat. Ross. 12: 142. 1916.
TYPE: Kazakhstan. Prov. Semirechinsk, Distr. Djar-
kinsk, Ketmen village, 5 July 1912, Saposhnikow &
Schischkin s.n. (lectotype, designated by Gonzalo et al.
[2011: 401], LE!); 5S. caucasica fo. brevifolia Roshev.,
Fl. Aziat. Ross. 12: 142. 1916. TYPE: Uzbekistan. Prov.
Fergana, Distr. Margelan. Kitchik-Alay River valley, at
1–2 km of Kindik mouth, 13 May 1914, Desiatoff 2186
(lectotype, designated by Gonzalo et al. [2011: 401],
LE!); 5S. caucasica fo. robusta Roshev., Fl. Aziat. Ross.
12:143.1916.TYPE:Kazakhstan.Prov.Syr-Darya,
Distr. Auliye-Ata, around Aleksandrovskaia village, 16
May 1909, Minkwitz 142 (lectotype, designated by Gon-
zalo et al. [2011: 401], LE!); S. caucasica var. typica
Drobow, Repert. Spec. Nov. Regni Veg. 21: 37. 1925.
nom. inval.; 5S. caucasica var. major Drobow, Repert.
Spec. Nov. Regni Veg. 21: 37. 1925. TYPE: not seen; 5
S. boczantzevii Tzvelev.
Habitat. Stipa caucasica var. caucasica is found in
steppes and rocky grasslands at 350–3000 m.
Distribution. Stipa caucasica var. caucasica is
widely distributed from Egypt through the Caucasus
up to central Asia (Pazij, 1968; Tzvelev, 1968, 1976;
Freitag, 1985; Wu & Phillips, 2006; Nobis, 2011;
Gonzalo et al., 2012, Nobis et al., 2019d). In Middle
Asia, it is found in Afghanistan, China, Iran, Kazakh-
stan, Kyrgyzstan, Tajikistan, Turkmenistan, and
Uzbekistan.
Stipa caucasica var. fanica M. Nobis, P. D. Gudkova
& A. Nowak, var. nov. TYPE: Kazakhstan. Steppe
grassland on hills & slopes, 6 km E of Targap
settlement, (80 km W of Almaty), 43°199220N/
75°559550E, alt. 775 m, exp. S, slope 5°, wp. 227,
18 May 2014, M. Nobis & P. Gudkova s.n. (holo-
type, KRA 486509!; isotypes, KRA 456384!, KRA
486503! KRA 486504!, KRA 486505!, KRA
486506!, KRA 486507!, KRA 486508!). Figure 6.
Diagnosis. Stipa caucasica Schmalh. var. fanica M. Nobis,
P. D. Gudkova & A. Nowak is similar to S. caucasica var.
caucasica in having a pilose lower segment of the awn (column)
with hairs 0.9–2(–2.5) mm long, but differs in having the
abaxial surface of vegetative leaves densely covered with
0.25–0.5 mm long hairs versus covered with ca. 0.1 mm long
prickles as well as in having the callus entirely pilose with
0.4–0.8 mm long hairs at the dorsal surface (ca. 2 times shorter
than those on the ventral surface) versus the callus entirely
pilose with hairs at the dorsal surface 1–1.5 mm long (com-
parable in length to those on the ventral surface), respectively.
Stipa caucasica var. fanica is also similar to S. drobovii var.
drobovii in having long hairs on the adaxial surface of the
leaves, but differs in having longer calluses (1.9–2.5 vs.
0.8–1.5 mm) with shorter hairs dorsally (0.4–0.8 vs.
1.5–3 mm).
Habitat. Stipa caucasica var. fanica is found in
steppes, stony slopes, and rocky grasslands at
350–2600 m.
Distribution. Stipa caucasica var. fanica is found
in the mountains of Middle Asia in Tajikistan (Turke-
stan, Zeravshan, western Pamir Mountains), Kyrgyzstan
(Fergana, Kirgiz, Tallas Mountains), Kazakhstan (Tian-
Shan Mountains), and Turkmenistan (Kopet-Dagh
Mountains).
Paratypes. KAZAKHSTAN. In N part of Balkhash Lake
near Gulshat settlement, 50 km SWW of Balkhash city, steppe
grassland, 46°419390N / 74°26960E, 360 m, 22 May 2014, M.
Nobis & P. Gudkova s.n. (KRA). KYRGYZSTAN. In the
eastern part of Tashkumyr town, 41°199470N / 72°129510E,
650 m, exp. S, slope 40°, no. 1, 11 May 2011, M. Nobis & A.
Nowak s.n. (KRA 479106); steppe grassland, ca. 3 km SE of
Kyzyl-Korgon village, 70 km SE of Osh, 40°99380N/
73°309410E, 6 June 2013, M. Nobis & A. Nowak 3/119
(KRA 479027); Karakabin region, 19 May 1952, Kirsanov
& Proskurnikov s.n. (KRA 455706); steppe grasslands in the
Talas River valley, WNW of Ivanovo-Alekseevka village near
Talas, alt. 1200 m, 42°339250N / 72°099320E, 10 May 2011,
M. Nobis s.n. (KRA 479055, KRA 479961); Adaevskii u.
Mangyshlak, Ak-tau, ur. Akmysh, 5 June 1926, M. D. Spi-
ridonov 693/2 (LE); Adaevskii u. Mangyshlak, okrestnostii
rodnika Dokarmysh, 4 June 1926, F. N. Rusanov 188/1
(LE); Turgai Distr., lower Sary-su River valley, Muyun-
kumov, near Kara-turgai, 31 May 1914, N. Krascheninnikov
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5/97 (LE); western Betpak-dala, Kol. Cholak-ecpe, 5 May
1936, B. Mironov & V. Pazij 712 (LE); Fergana Distr., Marge-
lanskii u., dol. r. Isfairat, Lyangar, 13 May 1913, N. Dessiatoff
1014 (LE); Karavanskii Distr., Dzhida-sai, 7 May 1952, Nabiev
71 (TASH). TAJIKISTAN. High mtn. steppes on the N slope of
Kuimond Mtns., in the Zeravshan River valley, to the S of
Langar settlement (ca. 80 km E of Aini), 39°249570N/
69°339160E, alt. 2000 m, exp. NNW, slope 30°, no. 3, 19
June 2009, M. Nobis s.n. (KRA 479052, KRA 479959, KRA
456385, KRA 479032, KRA 479035); Turkestan Mtns.,
Tashrabad, 1941, Pryakhin s.n. (TAD 1809); western Pamir,
S slope of Vakhan Mtns., alt. 2860 m, 3 June 1962, Nurban-
benov 134 (KRA 479960). TURKMENISTAN. Kopet-Dagh
Mtns., S of Prokhladnoe settlement, alt. 1450 m, 20 June
1934, A. G. Borisova 256 (TAD 1810).
Stipa caucasica subsp. nikolai M. Nobis, A. Nobis &
A. Nowak, subsp. nov. TYPE: Tajikistan. Fan
Mtns., high mtn. steppe grassland with share of
Juniperus,Rosa & other shrubs, ca. 1 km SE of
Sarytag settlement, on the right slope of Sarytag
stream valley [G´ory Fa ´nskie, wysokog´orska mur-
awa stepowa z udziałem Juniperus, Rosa i innych
krzew´ow, ok. 1 km na SE od wsi Sarytag, na prawym
zboczu doliny potoku Sarytag], 39°059380N/
68°169380E, alt. 2415 m., incl. NE, 15 June
2007, no. 4, M. Nobis, M. Kozak & A. Nowak s.n.
(holotype, KRA 454902!; isotypes, KRA 454912!,
KRA 454906!). Figure 6.
Diagnosis. Stipa caucasica Schmalh. subsp. nikolai M.
Nobis, A. Nobis & A. Nowak differs from S. caucasica subsp.
caucasica in having clearly shorter hairs in the lower and
middle part of the column (0.2–0.7[–0.9] vs. [0.9–]1–2[–2.5] mm
long). Additionally, the two subspecies differ in their habitat
preferences, with S. caucasica subsp. nikolai growing mainly in
upper mountain elevations and harsher habitats of rocky and stony
grasslands and steppes in Middle Asian mountains, whereas
S. caucasica subsp. caucasica prefers more arid habitats in low-
lands or lower mountain elevations in Middle Asia but also extends
further into the south and southwestern Asia.
Plants perennial, densely tufted, with a few culms
and numerous vegetative shoots; culms (20–)30–60 cm
tall, 3-noded, glabrous at nodes and very shortly pu-
bescent, or scabrous to glabrous below them. Leaves of
vegetative shoots: sheaths shortly pubescent, scabrous
to glabrous, at margins white and ciliate, cilia 0.5–1mm;
ligules rounded or truncate, 0–0.2 mm and ciliate
(0.4–1.5 mm) at margins; blades convolute, 5–35 cm
long, 0.5–0.8(–1) mm in diam., upper surface densely
short-pilose with hairs 0.09–0.12 mm, glabrous and
smooth beneath. Cauline leaves: sheaths glabrous to
slightly scabrous, rarely shortly pubescent in the upper
part, margins glabrous or ciliate, usually shorter than
internodes, upper sheath up to 20 cm, glabrous or sca-
brous, uninflated orslightly inflated; ligules 0.3–2.5 mm,
truncate to acute, ciliate at apex with hairs 0.3–1.5 mm
and setulose on the back; blades scabrous, up to 12 cm.
Panicle 8–25 cm, contracted, with 5 to 14 spikelets, at the
base enclosed by sheath of uppermost leaf, branches
erect, setulose, single or paired. Glumes subequal,
lower glume (33–)36–48(–60) mm, upper glume (32–)
35–46(–58) mm, narrowly lanceolate, tapering into long
hyaline apex, midvein sometimes setulose with cilia up to
1 mm. Anthecium (9–)10–12(–13.5) 30.8–1.1 mm. Callus
(1.7–)2–2.3(–2.6) mm, densely and long pilose ventrally,
with the longest hairs 1–1.9 mm, dorsally with straight hairs,
0.7–1.5 mm, base of callus not enlarged, peripheral ring
0.15–0.25 mm in diam., acute, cuneate, scar elliptic.
Lemma pale green, on dorsal surface with abundant
hooks and 7 lines of ascending hairs, 1–1.7 mm, ventral
lines terminating 0–3 mm below top of lemma, dorsal
line terminating (0–)0.5–4 mm below top of lemma; top
of lemma scabrous due to prickles and short hairs,
surpassed by a well-developed ring of unequal hairs
(0.3–)5–8(–1.2) mm at apex. Palea equal to lemma in
length, glabrous or with a dorsal line of ascending hairs
(hairs up to 0.6 mm), reaching up to 1/2–2/3 of palea
length. Awn (58–)85–110(–140) mm, unigeniculate or
slightly bigeniculate; column (14–)21–27(–36) mm,
twisted, 0.4–0.6 mm wide at base, shortly pilose, with
hairs (0.2–)0.3–0.7(–0.9) mm, in the upper part grad-
ually increasing in length toward second geniculation; seta
(40–)60–80(–110) mm long, falcate, (1.8–)2.5–3.3(–4.3)
times longer than column, hairs in lower part of seta (4–)
5–6(–7.3) mm, gradually decreasing in length toward apex.
Anthers purplish, 6–9 mm, glabrous.
Phenology. Stipa caucasica subsp. nikolai flowers
from May to June.
Habitat. Stipa caucasica subsp. nikolai is found in
stony and rocky grasslands and high mountain steppes
at 1000–3600 m.
Distribution. Stipa caucasica subsp. nikolai is a
widely distributed mountain taxon, occurring in Tian-
Shan, Pamir, Alai, and Karakorum Mountains. In Middle
Asia, it occurs in Afghanistan, Kazakhstan, Kyrgyzstan,
Tajikistan, and Uzbekistan.
Paratypes. KYRGYZSTAN. Western Tian-Shan, rocks
near the Sary-Chelek lake, 41°529N / 71°589E, alt. 1585 m,
9 June 2013, M. Nobis & A. Nowak s.n. (KRA 457808, KRA
457985, KRA 457986); Karasu River valley, Chatkalskii
regon, western Kyrgyzstan, western Tian-Shan, steppe grass-
lands in the Chat River valley (the left tributary of the Karasu
River), betw. Czat & Alcza settlement, 41°309320N/
72°149330E, alt. 1170 m, incl. SE, slope 30°, 10 May 2011,
M. Nobis s.n. (KRA 455307); ca. 5 km of Toktogul town,
41°519370N / 73°009530E, alt. 1020 m, 9 May 2011, M. Nobis
s.n. (KRA 455907); calcareous rocks, ca. 10 km N of Tash-
Kumyr town, 41°309320N / 72°149310E, alt. 1190 m, 9 June
2013, M. Nobis & A. Nowak s.n. (KRA 457548, KRA
457547). TAJIKISTAN. Zeravshan Mtns., Romit, on rocks,
5 June 2014, A. Nowak s.n. (KRA 455305, KRA 454417);
Zeravshan Mtns. high mtn. steppe ca 2.5 km E of Sarytag
settlement, 39°029380N / 68°199200E, alt. 2415 m, incl. S,
slope 5°, 15 June 2007, M. Nobis et al. s.n. (KRA 458310,
KRA 458311); Zeravshan Mtns. (Pamiro-Alai), high mtn.
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Nobis et al. 21
Synopsis of Stipa (Poaceae) in Middle Asia

steppe on the left slope of Fan River valley, near Zeravshan II
settlement, 39°129100N / 68°329080E, alt. 1610 m, incl.
SWW, slope 50°, 9 June 2009, M. Nobis s.n. (KRA
456371); Zeravshan Mtns., steppe on the left slope of Fan
River valley, Zeravshan settlement (S of Aini), 39°139080N/
68°319400E, alt. 1650 m, incl. NW, slope 25°, 10 June 2009,
M. Nobis s.n. (KRA 458510, KRA 458511); Jagnob River
valley, Zeravshan Mtns., rocks, left slope of the river valley,
ca. 1.5 km NW of Margeb settlement near Anzob, 39°129260N/
68°549410E, alt. 2220 m, incl. W, slope 80°, 9 June 2011,
M. Nobis s.n. (KRA 456372); Zeravshan Mtns., Kante River
valley,29June2014,A. Nowak s.n. (KRA 458320!); Zeravshan
Mtns., Remon settlement, alt. 2600 m, 18 June 2012, A. Nowak
s.n. (KRA 458323); Zeravshan Mtns. (Pamiro-Alai), steppe,
200 m S of tourists base Artucz, ca. 1 km S of Czukurak lake
(ca. 8 km SE of Madowrasettlement), 39°169140N / 68°089120E,
alt. 2225m, incl. NWW, slope35°, 25 June 2008, M. Nobis & M.
Kozak s.n. (KRA 454401, KRA 455288, KRA 454403, KRA
454404, KRA 455290); Zeravshan Mtns. steppe grassland near
the Artuch tourist camp, Artucz, ok. 2.5 km S of Czukurak lake
(ca. 8 km SE of Madowrasettlement), 39°169400N / 68°089210E,
alt. 2320 m, incl. S, slope 35°, 23 June2008, M. Nobis s.n. (KRA
454891, KRA 454426); Fan Mtns., stony steppe on the left slope
of Iskanderkul Lake, ca. 1.5–2 km SW of Varzob tourists camp,
39°049070N / 68°219050E, alt. 2220 m, incl. SE, 12 June 2007,
M. Nobis et al. s.n. (KRA 454409, KRA 454410, KRA 455294);
Mogiendaria River valley, steppe grassland, E of Khurmi set-
tlement near Mogien, 39°169280N / 67°419120E, alt. 1500 m,
exp. S, inc. 20° E, 12 June 2010, M. Nobis s.n. (KRA 455934);
Zeravshan Mtns., steppe grassland on the left slope of Mogien-
daria River valley, ca. 5 km NEE of Khurmi settlement near
Mogien, 39°169290N / 67°369100E, alt. 1520 m, exp. W-E, incl.
10–30° E, 27 May 2015, M. Nobis & A. Nowak s.n. (KRA
455906, KRA 455929); Zeravshan Mtns., Jagnob River valley,
steppe grassland, ca. 3–4 km E of Marzich settlement (W of
Anzob), 39°119210N / 68°449090E, alt. 2050 m, exp. S, incl.
30–70°, 22 June 2009, M. Nobis s.n. (KRA 455930), Pamir, high
mtn. steppe, in the Toguzbulok River valley(Szugnan Mtns.), ca.
8 km NW of Drzelondi settlement, 37°379520N / 72°329330E,
alt. 3520 m, 4 July 2008, M. Nobis s.n. (KRA 455933);
Zeravshan Mtns., Imat River valley, 8 km SW of Marguzor,
39°139050N / 68°219320E, 2650 m, incl. E, slope 35°, 22 June
2008, M. Nobis s.n. (KRA 455931, KRA 457974, KRA 457975,
KRA 457976, KRA 457977, KRA 457978, KRA 457979);
Zeravshan Mtns., Imat River valley, 9 km SW of Marguzor,
39°129520N / 68°219200E, alt. 2700 m, incl. SE, slope 35°, 22
June 2008, M. Nobis s.n. (KRA 451239, KRA 451241, KRA
451240, KRA 454896, KRA 451238, KRA 454895, KRA
451250, KRA 454908); Zeravshan Mtns. (Pamiro-Alai), high
mtn. steppe (in the Pasruddaria River valley) ca. 3 km W of
Marguzor settlement 39°159130N / 68°249040E, alt. 2190 m,
incl. S, slope10°, 19 June 2008, M. Nobis & A. Nowak s.n. (KRA
454415, KRA 454416, KRA 455293, KRA 454408); steppe
grasslands on the left slope of Iskanderkul Lake, ca. 500–1500 m
SW of tourist camp Varzob, 39°059050N / 68°229050E, 2220 m,
12 June 2007, M. Nobis et al. s.n. (KRA 457984, KRA 458313,
KRA 457982); Gabierud River valley, Pamiro-Alai,ca. 0.5 km S
of Shurpast settlement (S of Jagnob River), 39°079530N/
68°349520E, alt. 2300 m, incl. NE, slope 10° to 30°, 24 June
2009, M. Nobis et al. s.n. (KRA456370); Kazakhstan Chuilijskie
Mtns., near the Kurday pass, alt. 1100 m, 20 June 1978, L.
Ivaninia et al. s.n. (KRA 455308); Zeravshan Mtns., steppe, left
slope of the Czapdaria River valley, ca. 1 km NE of Allowodii
tourists camp (ca. 12 km NWW of Marguzor), 39°159460N/
68°169560E, alt. 2650 m, exp. SE, incl. 70°, 20 June 2008, M.
Nobis s.n. (KRA 455287, KRA 454400); Zeravshan Mtns., high
mtn. steppe on the left slope of Pasruddaria River valley, ca.
10 km NE of Allowodii touristscamp, 39°159080N / 68°199430E,
alt. 2460 m, exp. E, incl. 5°, 20 June 2008, M. Nobis s.n. (KRA
455291, KRA 454406, KRA 454405); Zeravshan Mtns., high
mtn. steppe on the left slope of Czapdaria River valley, ca. 1 km
NE of Allowodii tourists camp, 12 km NWW of Marguzor,
39°159460N / 68°169560E, alt. 2650 m, exp. SE, incl. 70°,
20 June 2008, M. Nobis s.n. (KRA 455283, KRA 454398,
KRA 455282, KRA 455281); Zeravshan Mtns., high mtn. steppe
in the eastern part of Marguzor settlement (near the rd.),
39°139530N / 68°269020E, alt. 2246 m, exp. NW, slope 50°, 19
June 2008, M. Nobis s.n. (KRA 455292, KRA 454407); high
mtn. steppe, on the left slope of Uriecz settlement, ca. 1 km NW
of Kulikalon Lake (Gowkhona Mtns.) (ca. 8 km SE of Madowra
settlement), 39°159560N / 68°109140E, alt. 2750 m, exp. NE,
slope 25°, 25 June 2008, M. Nobis & M. Kozak s.n. (KRA
455289, KRA 454402); steppe grassland on the left slope of
Uriecz River valley, ca. 4 km S of Jakahona settlement (2 km N
Artucz tourists camp), 39°179040N / 68°079360E, alt. 2240 m,
exp. SW, slope 30°, 26 June 2008, M. Nobis s.n. (KRA 454884,
KRA 454423); Zerawszan Mtns., high mtn. steppe on the left
slope of Pasruddaria River valley, ca. 3 km NE of Allowodii
tourists camp, 39°159240N / 68°189530E, alt. 2550 m, exp. S,
slope 30°, 20 June 2008, M. Nobis s.n. (KRA 455285, KRA
454399); Zeravshan Mtns., high mtn. steppe, betw. stones, on
the left slope of Iskanderdarya River valley, ca. 0.5 km E of
Serimadarun Lake (near Iskanderkul Lake), 39°059080N/
68°229460E, alt. 2326 m, incl. W, slope 5° to 10°, 14 June 2011,
M. Nobis s.n. (KRA 455905); Fann Mtns., high mtn. steppe on
the left slope of Sarytag stream valley, 39°039080N/
68°199010E, alt. 2400 m, incl. S, 16 June 2008, M. Nobis s.n.
(KRA 454897, KRA 451242); Fann Mtns., high mtn. steppe on
the left slope of Arch stream valley, NW of Sarytag settlement,
39°039070N / 68°199170E, alt. 2410 m, incl. SW, 16 June 2008,
M. Nobis s.n. (KRA 451243, KRA 451244, KRA 451245, KRA
451246, KRA 451247, KRA 454899); Fan Mtns., steppe grass-
lands on the right slope of Karakul River valley, 3–4kmSWof
Sarytag settlement, 39°029090N / 68°169450E, alt. 2460 m, incl.
NW, 17 June 2008, M. Nobis s.n. (KRA 454412, KRA 455300,
KRA 454411); Zeravshan Mtns., stony steppe on the right slope
of Pasruddaria River valley, ca. 6 km NE of Marguzor settlement
39°149570N / 68°229150E, alt. 2350 m, exp. N, slope 15° to 25°,
22 June 2008, M. Nobis s.n. (KRA 454904, KRA 451249, KRA
454903, KRA 451248); Fan River valley, Gushty Mtns., ca.
500 m E of Zeravshan settlement, steppe, 39°139140N/
68°329510E, alt. 2150 m, exp. W, slope 25°, 21 June 2009,
M. Nobis & A. Nowak s.n. (KRA 457922, KRA 458324,
KRA 458325, KRA 458327).
Stipa chitralensis Bor, Kew Bull. 9: 500. 1954. TYPE:
Pakistan. Chitral, Guger, 18 May 1895, Harris
16800 (holotype, K!; isotypes, BM!, E!, WU not
seen). [Neotrinia chitralensis (Bor) M. Nobis,
comb. nov.
Notes. Contrasting with other species of Stipa,spec-
imens representing Neotrinia chitralensis have 2–2.5 mm
long hyaline lobes on the apical part of the lemma. Rep-
resentatives of Stipa have lemmas without elongated lobes
or lobes (if present) are flat and less than 1 mm long (Freitag,
1985). In N. chitralensis, the pattern of its lemma micro-
morphology is dominated by elongated basal cells and
frequent silica bodies and cork cells (M. Nobis, pers.
obs.), which makes it more similar to N. splendens (Trin.)
M. Nobis, P. D. Gudkova & A. Nowak rather than other
22 Annals of the
Missouri Botanical Garden

Figure 7. Patterns of the lemma micromorphology in selected species of the tribe Stipeae: (a) Achnatherum haussknechtii
(Boiss.) M. Nobis (Iran, J. Bornm ¨uller 4834 [PRC]), (b) A. staintonii (Bor) M. Nobis & P. D. Gudkova (Nepal, M. A. Farille 81-340
[E]), (c) A. mandavillei (Freitag) M. Nobis (Oman, Mandaville 6525 [KAS]), (d) A. turkomanicum (Roshev.) Tzvelev (Tajikistan,
M. Nobis s.n. [KRA]), (e) A. calamagrostis (L.) Beauv. (Hungary, H. Petry s.n. [KRA]), (f) A. caragana (Trin.) Nevski (Tajikistan,
M. Nobis s.n. [KRA]), (g) A. pubicalyx (Ohwi) Keng (China, Kozlov 124 [LE]), (h) A. inebrians (Hance) Keng ex Tzvelev (China,
M. Olonova s.n. [KRA]), (i) A. bromoides (L.) P. Beauv. (Iran, T. Alexeenko 316 [LE]), (j) A. brandisii (Mez) Z. L. Wu (India, Kashmir,
R. R. Stewart 18120 [NY]), (k) Stipellula capensis (Thunb.) R¨oser & Hamasha (Spain, R. Piwowarczyk s.n. [KRA]), (l) Ptilagrostis
malyschevii Tzvel. (Kyrgyzstan, M. Nobis s.n. [KRA]), (m) P. alpina (Fr. Schmidt) Sipl. (Russia, S. Kharkevich, T. Bush s.n. [NY]),
(n) P. mongholica (Turcz. ex Trin.) Griseb. (Mongolia, A. Pacyna s.n. [KRA]), (o) Orthoraphium roylei (Nepal, M. A. Farille s.n. [E]),
(p) Macrochloa tanacissima (L.) Kunth (Spain, R. Piwowarczyk s.n. [KRA]), (r) Neotrinia splendens (Trin.) M. Nobis, P. D. Gudkova
& A. Nowak (Tajikistan, Yu. Gusev s.n. [LE]), (s) Piptatherum munroi (Stapf ex Hook. f.) Mez (Nepal, M. F. Watson et al. DNEP3
AX33 [E]), (t) Patis coreana (Honda) Ohwi (China [KUN 387221]), (u) Stipa 3brevicallosa M. Nobis (Tajikistan, M. Nobis s.n.
[KRA]), (v) S. grandis P.A. Smirn. (Mongolia, Safronova et al. s.n. [KRA]), (w) S. bungeana Trin. (Kyrgyzstan, M. Nobis & A. Nowak
s.n. [KRA]), (x) S. drobovii (Tzvelev) Czerep. (Tajikistan, M. Nobis s.n. [KRA]), (y) S. zeravshanica M. Nobis (Tajikistan, M. Nobis
s.n. [KRA]). Abbreviations: l, long cell (fundamental cell); s, silica cell (silica body); c, cork cell; h, hook; mh, macrohair.
Volume 105, Number 1
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Nobis et al. 23
Synopsis of Stipa (Poaceae) in Middle Asia

members of Stipa or Achnatherum in which the lemma
epidermal pattern is different (Fig. 7; see also Romaschenko
et al., 2012; Nobis et al., 2019a, 2019b). Thus, the transfer
of S. chitralensis to the genus Neotrinia is proposed.
Stipa confusa Litv., Izv. Akad. Nauk SSSR, Ser. 6, 1928:
53, pl. 3, f. 2. 1928. TYPE: Altai, ad fluvium
Tscharysch in montosis herbidis, July 1826, Ledebour
172 (lectotype, designated by Tzvelev [1976: 576],
LE TRIN 1441.2!; isolectotype, LE TRIN 1441.4!).
[Achnatherum confusum (Litv.) Tzvelev, Probl.
Ekol. Geobot. Bot. Geogr. Florist. 140. 1977.
Stipa 3consanguinea Trin. & Rupr., Sp. Gram.
Stipac. 78. 1842, pro sp. (S. krylovii Rochev. 3
S. glareosa P. A. Smirn.). TYPE: Altai, desert
Tschuj. edit., 1833, Dr. Bunge s.n. (lectotype,
designated here, LE Herbarium Trinii 1393.1!;
isolectotypes, LE! [2 sheets]).
Habitat. Stipa 3consanguinea is found in steppe
grasslands and high mountain deserts at 1800–3000 m.
Distribution. Stipa 3consanguinea is found in
mountains of north-central Asia (Tian-Shan: China,
Altai: Russia, Mongolia, Kazakhstan; Tzvelev, 1968,
1976). In Middle Asia, it occurs in Kazakhstan and
western China (Tian-Shan Mountains).
Notes. Stipa 3consanguinea was described by Tri-
nius and Ruprecht (1842) from Altai. The original
description states: “Fl. Altalca: in desertis Tschujae
editioribus; Julio (Bunge!).”The collection of four
syntypes of this taxon is preserved at LE. The first
typification of this species was provided by Tzvelev
(1976: 580), who instead of choosing a lectotype indi-
cated that a [holo-] type and two isotypes were preserved
at LE. Later, Byalt (in Sokolova, 2012: 300), citing
Tzvelev (1976), designated the lectotype, isolectotype,
and two syntypes and attached appropriate labels to the
sheets in 2010. Unfortunately, this lectotypification of
S.3consanguinea is not effective because he did not use the
phrase “designated here”(see Art. 7.11 of the International
Code of Nomenclature, McNeill et al., 2006, 2012; Turland
et al., 2018). Because Byalt in Sokolova (2012) did not use
the required phrase, we provide the correct designation of the
lectotype. We designated specimens on the herbarium sheet
preserved in the Trinius herbarium and deposited them at LE
as the lectotype; these were previously designated as the
holotype by P. Peterson (Soreng et al., 1995) because infor-
mation on the herbarium label is most similar to that in the
original description. The other two sheets with labels “Herb.
Fisch., in lapidosis apricis ad Tschuja, [18]32, Bunge (LE)”
and “Stipa juncea ?, Bge fl alt suppl., in lapidosis apricis ad
fluv. Tschuja, 1832, Dr. Bunge, Acc. ab inventore 1833,
S. consanguinea Rupr. Hb. Meyer (LE 01011394)”are
isolectotypes. The fourth sheet, which has one specimen
and a label reading “Herb. Fisch. Stipa consanguinea Trin.,
Stipa juncea L. ?, suppl. fl. alt., Bge.,”is a syntype.
Stipa crassiculmis P. A. Smirn., Repert. Spec. Nov.
Regni Veg. 22: 375. 1926. TYPE: Kopet-Dagh,
pereval Arvaz, 2300 m, Kultiasov s.n. (holotype,
TASH not seen). [Stipa pulcherrima K. Koch
subsp. crassiculmis (P. A. Smirn.) Tzvelev.
Stipa 3czerepanovii Kotukhov, Turczaninowia 1(2):
13. 1998 (S. orientalis Trin. 3S. richteriana Kar.
& Kir.; Nobis & Gudkova, 2016). TYPE: [Kazakh-
stan.] Depressio Zaissanica denudatinies argillarum
tertiariarum Akseir (maryo orientalis) jugi parve decli-
vitas australi-occidentalis glareoso-argillosa, 17 July
1993, Ju. Kotuchov s.n. (lectotype, designated by Nobis
& Gudkova [2011: 199], LE!; isolectotypes, KRA
436047!, KRA 436048!, KUZ! [9 sheets], LE!).
Habitat. Stipa 3czerepanovii is found in steppe
grasslands at 600–1500 m.
Distribution. Stipa 3czerepanovii is found in east-
ern Kazakhstan (Kotukhov, 2002; Nobis & Gudkova,
2016) (Fig. 4).
Stipa damascena Boiss., Diagn. Pl. Orient. ser. 1, 13:
45–46. 1854. TYPE: [Syria.] In collibus aridissi-
mis pone urbem Damascum, May 1846, Boissier
(holotype, G not seen; isotype, W!). 5Stipa arab-
ica Trin. & Rupr. var. turgaica (Roshev.) Tzvelev.
Stipa dasyphylla (Lindem.) Trautv., Trudy Imp. S.-
Peterburgsk. Bot. Sada 9: 350. 1884. TYPE:
Kharkov, na kholmakh i kosogorakh Rogani, 13
June 1853, V. Chernyaev s.n. (lectotype, designated
by Tzvelev [1976: 589], LE!; isolectotype, LE!).
Habitat. Stipa dasyphylla is found in steppe grass-
lands at 1500–1600 m.
Distribution. Stipa dasyphylla is found from eastern
Europe to northern and central Asia (Tzvelev, 1968,
1976; Kotukhov, 2002). In Middle Asia, it occurs in
Kazakhstan (Manrak Mountain).
Stipa decipiens P. A. Smirn., Ucen. Zap. Moskovsk.
Gosud. Univ. 2: 338. 1934. TYPE: Altai merid., in
declivibus stepposis, locis lapidosis prope pag.
Katon-Karagaj, ca. 1100 m, 24 June 1930, P. Smirnov
3a (holotype, MW!; isotype, MW!). 5Stipa krylovii
Roshev.
Stipa densa P. A. Smirn., Del. Sem. Hort. Bot. Univ.
Mosq. 15, 1930 (replaced synonym S. densiflora
P. A. Smirn., nom. illeg.). TYPE: [Russia.] Kha-
kasia, okr. sel. Askyz, slabo solontsevataya khrya-
schevataya step’v doline Abakana, July 1927, V. V.
Reverdatto s.n. (holotype, TK!). 5Stipa krylovii
Roshev.
24 Annals of the
Missouri Botanical Garden

Stipa densiflora P. A. Smirn., Repert. Spec. Nov. Regni
Veg. 26: 265. 1929, nom. illeg. non Hughes 1921.
TYPE: [Russia.] Khakasia, okr. sel. Askyz, slabo solo-
ntsevataya khryaschevataya step’v doline Abakana,
July 1927, V. V. Reverdatto s.n. (holotype, TK!). 5
Stipa krylovii Roshev.
Stipa desertorum (Roshev.) Ikonn., Opred. Vyssh. Rast.
Badakhshana 83. 1979. Basionym: Stipa caucasica
Schmalh. fo. desertorum Roshev., Fl. Aziat. Ross.
12: 143. 1916. TYPE: Tsentr. Tian-Shan, Przhe-
valskii u., u Tamchi bliz oz. Issyk-Kul’, na peskakh,
19 June 1908, R. Rozhevich 574 (lectotype, desig-
nated by Tzvelev [1976: 593], LE!). 5Stipa glar-
eosa P. A. Smirn. var. glareosa.
Stipa drobovii (Tzvelev) Czerep., Sosud. Rast. SSSR
387. 1981 [replaced synonym: S. bella Drobow,
hom. illeg.]. TYPE: Uvaly Karatau severnoe oz.
Biilikol’, slantsevye sklony k oz. Akkul’, 1 June
1922, V. Drobov 285 (lectotype, designated by
Tzvelev [1976: 592], LE!; isolectotype, TASH831!).
Notes. Having a short callus with very distinct, long,
and falcate hairs dorsally, Stipa drobovii is an easily
distinguishable taxon, but varies considerably in the
pubescence of its leaves (blades and sheaths). Based on
the revised plant material, four varieties are recognized
in Middle Asia.
1. Abaxial surface of leaves of vegetative shoots densely
pubescent...............Stipa drobovii (Tzvelev)
Czerep. var. iskanderkulica (Tzvelev) M. Nobis & A. Nowak
19. Abaxial surface of leaves of vegetative shoots gla-
brous and smooth . . . . ..........................2
2. Adaxial surface of vegetative leaves covered with
hairs up to 0.1 mm long . . . . . . . . . . . . . . . . . . . . .
Stipa drobovii var. persicorum M. Nobis
29. Adaxial surface of vegetative leaves covered with
hairs 0.3–0.6 mm long . . ........................3
3. Sheaths of cauline leaves shortly and densely pu-
bescent . . . . .....Stipa drobovii var. jarmica M. Nobis
39. Sheaths of cauline leaves glabrous . . . . . . . . . . . . .
........................ Stipa drobovii var. drobovii
Stipa drobovii var. drobovii.
Habitat. Stipa drobovii var. drobovii is found in
steppe grasslands at 800–2500 m.
Distribution. Stipa drobovii var. drobovii is widely
distributed from the Caucasus to Middle Asia (Pazij,
1968; Tzvelev, 1968, 1976; Freitag, 1985; Wu & Phillips,
2006; Gonzalo et al., 2012). In Middle Asia, it occurs in
Afghanistan, Iran, Kazakhstan, Kyrgyzstan, Tajikistan,
Turkmenistan, and Uzbekistan.
Stipa drobovii var. iskanderkulica (Tzvelev) M. Nobis
& A. Nowak, Phytotaxa 303(2): 151. 2017. Basio-
nym: Stipa caucasica Schmalh. subsp. iskanderkul-
ica Tzvelev, Novosti Sist. Vyssh. Rast. 11: 20. 1974.
TYPE: Samarkand Distr., Iskanderkul, 21 May
1914, V. Dubyanskii s.n. (holotype, LE!).
Synonyms. [Stipa iskanderkulica (Tzvelev) Czerep.;
5S. caucasica Schmalh. f. dasyphylla Roshev., Fl. Aziat.
Ross. 12: 142. 1916. TYPE: LE not seen; 5S. bella
Drobow var. incana Korol. ex Ovcz. & Czukavina, Fl.
Tadzhik. 1: 421. 1957, nom. nud.
Habitat. Stipa drobovii var. iskanderkulica is found
in steppe grasslands at 1500–2500 m.
Distribution. Stipa drobovii var. iskanderkulica is
found in Middle Asia (Ovchinnikov & Chukavina,
1957; Nobis et al., 2017a), specifically in Tajikistan
(Zeravshan and Hissar Mountains).
Stipa drobovii var. jarmica M. Nobis, var. nov. TYPE:
Dolina rzeki Zeravshan (Zeravshan River valley),
Rejon Zerawsza ´nski C (Zeravshan Region C), G ´ory
Zeravsha ´nskie (Pamiro-Alai); Murawy stepowe na
przydro˙znych skarpach, na prawej terasie rzeki
Zerawszan, w pobli˙zu wsi Diemnora, w pobli˙zu
potoku Liangarshif (około 100 km na E od Aini)
[near Demnora settlement, near Langarshif stream
(ca. 100 km E of Aini), steppe grasslands on the
roadside escarpment, on the right terrace of Zer-
avshan River], 39°279000N / 69°529360E, alt. 2450 m,
exp. S, slope 30°, loc. 7, 19 June 2009, M. Nobis s.n.
(holotype, KRA 457708!; isotypes, KRA 454144!,
KRA 456642!, KRA 456646!, KRA 457709!).
Diagnosis. Stipa drobovii (Tzvelev) Czerep. var. jarmica M.
Nobis resembles S. drobovii var. drobovii in having vegetative
leaves that are glabrous on the abaxial surface but differs from
the latter taxon in having cauline leaves with densely pubescent
(vs. glabrous) sheaths.
Plants perennial, densely tufted, with a few culms
and numerous vegetative shoots; culms (35–)40–60 cm
tall, 3-noded, glabrous at nodes and densely pubescent
below them. Culm leaves: lower, middle, and upper
sheaths densely shortly pubescent, blades glabrous and
smooth on abaxial surface and densely pubescent on
adaxial surface. Leaves of the vegetative shoots: ligules
up to 0.2 mm, densely ciliate, cilia up to 1.5 mm; abaxial
surface of blades glabrous and smooth, whereas adaxial
surface densely covered with hairs 0.25–0.6 mm.
Glumes 32–46 mm. Anthecium 8–11 30.8–1.1 mm.
Callus acute, 0.8–1.4 mm, its base 0.6–0.9 3
0.25–0.45 mm, densely bearded, dorsal hairs falcate
and longer than ventral hairs. Lemma with 5 to 7 lines of
hairs, marginal and dorsal lines reaching base of awn or
terminating at 0.2–1 mm below top; lemma apex with
ring of hairs. Awn unigeniculate, column 12–20 mm;
with hairs 13–20 mm; seta arcuate, 40–65 mm, hairs in
lower part of seta (35–)40–50 mm.
Volume 105, Number 1
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Nobis et al. 25
Synopsis of Stipa (Poaceae) in Middle Asia

Phenology. Stipa drobovii var. jarmica flowers from
May to June.
Habitat. Stipa drobovii var. jarmica is found in
steppe grasslands at 1900–2500 m.
Distribution. Stipa drobovii var. jarmica is found in
Tajikistan (Turkestan and Zeravshan Mountains).
Etymology. The name of the taxon originates from
the name of the Jarm (Yarm) settlement located in the
eastern part of the Zeravshan River valley.
Paratypes. TAJIKISTAN. Northern slope of Turkestan
Mtns., on the left bank of Argly River valley, foreland of the
Kurganak Mtn., 24 July 1967, A. N. Konnov 2542 (KRA); high
mtn. steppes on the left slope of Kuimont Mtns., in the Zeravshan
River valley, to the S of Langar settlement (ca. 80 km E of Aini),
39°249570N / 69°339160E, alt. 2000 m, exp. NNW, slope 30°,
loc. 3, 19 June 2009, M. Nobis s.n. (KRA 455387, KRA 455808,
KRA 455811,KRA 455812, KRA 455850, KRA 456212, KRA
456636, KRA 457706, KRA 457443, KRA 456650); high mtn.
steppes on the left slope of Kuimont Mtns., at the mouth of the
river Tabaspin to the Zeravshan, ca. 2 km S of Langar settlement
(ca.80kmEofAini),alt.2340m,exp.W,slope50°,loc.2,19
June 2009, M. Nobis s.n. (KRA 457428, KRA 457444, KRA
457159, KRA 455343); steppe grassland on the right terrace of
Zeravshan River near the Sabakh settlement, near the mouth of
Dzhikhdon River (ca. 100 km E of Aini), alt. 2350 m,
39°289350N / 69°479290E, exp. S, slope 10° to 30°, loc. 2,
19 June 2009, M. Nobis s.n. (KRA 456637, KRA 456638, KRA
456639, KRA 456640, KRA 456641, KRA 455809, KRA
455810, KRA 456644, KRA 456645, KRA 457707); steppe
grassland on the left slope of Zeravshan River valley (near the
rd.), ca. 4 km E of Tabusht settlement, W of Yarm (ca. 115 km E
of Aini), alt. 2360 m, 39°279470N / 69°479520E, exp. SE, slope
10° to 30°, loc. 5, 20 June 2009, M. Nobis s.n. (KRA 457429);
steppe grassland on the right terrace of Zeravshan River valley,
ca. 3–4 km W of Yarm settlement (ca. 125 km E of Aini), alt.
2455 m, 39°269100N / 69°569040E,exp. S, slope 10° to 60°, loc.
4, 20 June 2009, M. Nobis s.n. (KRA 457431, KRA 457433,
KRA 453742, KRA 453743).
Stipa drobovii var. persicorum M. Nobis, var. nov.
TYPE: Persia [Iran]. Ad pedem merid, montium
K´uh-e Shan Jahan, ca. 20 km austro-orient. ab
oppido Esfarain, 36°579–57°409, 16 June 1977,
J. Soj ´ak s.n. (holotype, PR!).
Diagnosis. Stipa drobovii (Tzvelev) Czerep. var. persicorum
M. Nobis is similar to S. drobovii var. drobovii in its lower segment
of the awn (column) long pilose, with hairs 0.9–2(–2.5) mm, but
differs in having the abaxial surface of the vegetative leaves
densely covered with hairs 0.10–0.15 mm, whereas S. drobovii
var. drobovii has leaves covered with hairs 0.25–0.5 mm.
Plants perennial, densely tufted, with a few culms
and numerous vegetative shoots; culms (15–)20–30 cm
tall, 3-noded, glabrous at nodes and densely pubescent
below them. Culm leaves: sheaths glabrous, blades
glabrous and smooth on abaxial surface and pubescent
on adaxial surface. Leaves of vegetative shoots: ligules
up to 0.2 mm, densely ciliate, cilia up to 1.5 mm; abaxial
surface of blades glabrous and smooth, whereas densely
covered with hairs 0.1–0.15 mm adaxially. Glumes
34–45 mm. Anthecium 8–10 30.8–1.0 mm. Callus
acute, 0.8–1.2 mm, its base 0.6–0.9 30.25–0.45 mm,
densely bearded, dorsal hairs falcate and longer than
ventral hairs. Lemma with 5 to 7 lines of hairs, marginal
and dorsal lines reaching base of awn or terminating at
0.2–1 mm below top; lemma apex with ring of hairs.
Awn unigeniculate, column 12–20 mm; with hairs
13–20 mm; seta arcuate, 40–65 mm, hairs in lower
part of seta (35–)40–50 mm.
Phenology. Stipa drobovii var. persicorum flowers
from April to June.
Habitat. Stipa drobovii var. persicorum is found in
steppes, stony slopes, and rocky grasslands at
1500–2000 m.
Distribution. Stipa drobovii var. persicorum is found
in Middle Asia, specifically in northeastern Iran and
Turkmenistan (Kopet-Dagh Mountains).
Stipa duthiei Hook. f., Fl. Brit. India 7: 232. 1896.
TYPE: [India.] Tehri Garwhal, Lekhus, below
Srikanta, 12,000–13,000 ft., 11 Aug. 1853, Duthie
273 (holotype, K 32097!). [Ptilagrostis duthiei
(Hook. f.) M. Nobis & P. D. Gudkova, PhytoKeys
128: 107. 2019 [[Achnatherum duthiei (Hook. f.)
P. C. Kuo & S. L. Lu, Fl. Reipubl. Popularis Sin.
9(3): 322, pl. 80, f. 9–14. 1987].
Stipa 3dzungarica M. Nobis, nothosp. nov. (S. kirghi-
sorum P. A. Smirn. 3S. richteriana Kar. & Kir.
subsp. richteriana). TYPE: Kazakhstan. Western part
of Dzhungarskii Alatau, N Tian-Shan, ca. 3 km NW
of Sary-Ozek, ca. 140 km NNE of Almaty, steppe,
44°229580N / 77°569160E, 1007 m, wp. 1250, 27
May 2019, M. Nobis (holotype, KRA 479043!; iso-
type, KRA!).
Diagnosis. Stipa 3dzungarica M. Nobis is similar to
S. 3heptapotamica Golosk., but it differs from the latter taxon
in having longer anthecia, 9.5–11.5 mm (vs. 7–9.2 mm), with
hairs arranged in 7 lines and without a ring of hairs at the apex
(vs. anthecia pilose all over with a well-developed ring of hairs
at the apex), longer calluses, 1.5–2.2 mm (vs. 1.3–1.6 mm), and
longer glumes, 18–28 mm (vs. 15–20 mm).
Plants perennial, densely tufted, with a few culms
and numerous vegetative shoots; culms 50–58 cm tall,
3-noded, glabrous at nodes and very shortly pubescent
below them. Leaves of vegetative shoots: sheaths shortly
pubescent; ligules rounded or truncate, up to 0.2 mm
and ciliate at margins; blades convolute, up to 35 cm,
0.4–0.6(–0.7) mm in diam., adaxial surface densely
pubescent with hairs 0.1–0.15 mm, abaxial surface
scabrous due to hooks and short prickles. Cauline
leaves: sheaths glabrous and with white edges, shorter
than internodes, glabrous; ligules 0.2–0.5 mm, obtuse
26 Annals of the
Missouri Botanical Garden

and ciliate at margins; blades scabrous, up to 8 cm.
Panicle 18–25 cm, contracted, at base enclosed by
sheath of uppermost leaf, branches erect, setulose,
single or paired. Glumes subequal, 18–28 mm, narrowly
lanceolate, tapering into long hyaline apex, midvein
sometimes setulose. Anthecium 9.5–11.5 30.7–1 mm.
Callus 1.5–2.2 mm, densely long-pilose on ventral and
dorsal surfaces, callus base not enlarged, peripheral
ring 0.2 mm in diam., acute, cuneate, scar elliptic to
circular. Lemma pale green, on dorsal surface with
abundant hooks and with 7 lines of ascending hairs,
hairs up to 0.6 mm, ventral line of hairs terminating at
0.4–1 mm below top of lemma and dorsal line termi-
nating at 2–3 mm below top of lemma; top of lemma
scabrous due to hooks and prickles but without ring of
hairs at apex. Palea equal to lemma in length. Awn
125–155 mm, bigeniculate; lower segment of column
34–47 mm, twisted, ca. 0.5 mm wide near base, dis-
tinctly scabrous due to hard prickles, gradually increas-
ing in length toward geniculation; upper segment of
column 12–15 mm, twisted, scabrous due to prickles
and short hairs 0.1–0.3 mm, gradually increasing in
length toward geniculation; seta straight or flexuous
90–120 mm, hairs in lower part of seta 1.2–1.5(–1.7)
mm, gradually decreasing in length toward apex. An-
thers yellow, 4–5 mm, glabrous.
Phenology. Stipa 3dzungarica flowers from May to
June.
Habitat. Stipa 3dzungarica is found in steppes at
800–1100 m.
Distribution. Stipa 3dzungarica is found in Middle
Asia, specifically Kazakhstan (in the western part of
Dzhungarskii Alatau).
Notes. The plot of vegetation that includes Stipa 3
dzungarica is documented here: feather grass steppe,
on the loess substrate, area of relev´e16m
2
, cover of
herbs layer 70%: S. 3dzungarica 1,S. kirghisorum 3,
S. richteriana subsp. richteriana 1, S. sareptana 2,
Bromus tectorum 1, Artemisia sp. 3, Ziziphora tenuior 1,
Poa bulbosa 1, Salvia sp. 1,Holosteum glutinosum 1,
Orobanche amoena 1,Iris cf. glaucescens 1,Descurainia
sophia 1,Festuca valesiaca 1. Steppes in the southwest part
of Dzungarskii Alatau are rich in species of feather grasses.
At the locality near Sary-Ozek, together with S. 3dzungar-
ica, the following taxa were recorded: S. 3heptapotamica,
S. kirghisorum,S. sareptana,S. orientalis,S. 3czerepanovii,
S. richteriana subsp. richteriana,S. arabica,andS. cauca-
sica. It is possible that further localities of S. 3dzungarica
can be found in the region.
Stipa effusa Mez, Repert. Spec. Nov. Regni Veg. 17:
210. 1921. TYPE: Persia austro-orient., Prov.
Kerman, Kuh-e-Dschupar, in reg. alp. et subalp.,
2900–3200 m, 9 June 1892, Bornmuller 4837
(lectotype, designated here, JE!; holotype, pre-
served at B, was destroyed during World War
II). 5Achnatherum haussknechtii (Boiss.)
M. Nobis [[Stipa haussknechtii Boiss.].
Stipa 3fallax M. Nobis & A. Nowak, Phytotaxa 303(2):
145. 2017 (S. drobovii (Tzvelev) Czerep. 3S.
macroglossa P. A. Smirn. subsp. macroglossa; Nobis
et al., 2017a). TYPE: Western Tajikistan, western
Pamir Alai. Zeravshan Mtns., high mtn. steppe, on
the southern slope of mtns. (left slope of the Iskan-
derdarya River valley), ca. 0.6 km E of Serimadarun
Lake (near Iskanderkul Lake), 39°059N / 68°239E,
alt. 2340 m, incl. S, slope 5°, 15 June 2012, M.
Nobis s.n. (holotype, KRA 407904!; isotypes, KRA
407898!, KRA 407899!, KRA 407900!, KRA
407901!, KRA 407902!, Herb. Stip. M. Nobis!).
Habitat. Stipa 3fallax is found in mountain steppe
grasslands at 2200–2400 m.
Distribution. Stipa 3fallax is found in Middle Asia
(Nobis et al., 2017a), specifically in Tajikistan (Zerav-
shan Mountains). Figure 3.
Stipa glareosa P. A. Smirn., Repert. Spec. Nov. Regni
Veg. 26: 266. 1929. TYPE: Gobi. Steppum glar-
eosum in depressione lac. Orok-nor, 7 Sep. 1924,
N. Pavlov 169 (holotype, MW!; isotype, LE!).
Synonyms. [Stipa caucasica Schmalh. subsp.
glareosa (P. A. Smirn.) Tzvelev, Novosti Sist. Vyssh.
Rast. 11: 20. 1974; 5Stipa orientalis Trin. var. tricho-
glossa Hack., Vidensk. Meddel. Dansk Naturhist.
Foren. Kjøbenhavn 55: 164. 1903. TYPE: Ad Sufi
Kurgan in montibus Alai, 18 July 1898, Paulsen 404
(lectotype, designated by Gonzalo et al. [2011: 405], C;
isolectotype, W!); 5S. glareosa var. langshanica Y. Z.
Zhao, Acta Sci. Nat. Univ. Nei Menggu 23(4): 546.
1992. TYPE: China. Langshan, 4 July 1988, Zhao Yizhi
et al. 4077 (holotype, HIMC not seen); [S. langshanica
(Y. Z. Zhao) Y. Z. Zhao; 5S. caucasica Schmalh. fo.
desertorum Roshev., Fl. Aziat. Ross. 12: 143. 1916.
TYPE: Tsentralnyi Tyan-Shan, Semirechinskaya oblast,
Przhevalskii uezd, u reki Tamachi, bliz oz. Issyk-kul, na
peskakh, 19 June 1908, Roshevitz 574 (lectotype, des-
ignated by Tzvelev [1974: 20], LE!); [S. caucasica var.
desertorum (Roshev.) Tzvelev in V. I. Grubov, Pl. Cen-
tral Asia 4: 53. 1968; [S. caucasica subsp. desertorum
(Roshev.) Tzvelev, Novosti Sist. Vyssh. Rast. 11: 20.
1974; [S. desertorum (Roshev.) Ikonn.
Habitat. Stipa glareosa is found in gravelly or sandy
deserts, steppes, high mountain rocky grasslands, and
screes at 350–5000 m.
Volume 105, Number 1
2020
Nobis et al. 27
Synopsis of Stipa (Poaceae) in Middle Asia

Distribution. Stipa glareosa is a widely distributed
central Asian species (Pazij, 1968; Tzvelev, 1968, 1976;
Freitag, 1985; Dickor´e, 1995; Wu & Phillips, 2006;
Gonzalo et al., 2012; Gudkova et al., 2013a, 2015; Nobis,
2014). It also occurs in Middle Asia in Afghanistan,
China, Kazakhstan, Kyrgyzstan, and Tajikistan.
Notes. Stipa caucasica fo. desertorum differs from S.
glareosa in having glabrous and smooth versus scabrous
vegetative leaves (Roshevitz, 1916; Gudkova et al.,
2013b). However, during field studies, specimens with
both glabrous and/or slightly scabrous leaves were ob-
served and collected by us from localities where S. glareosa
was recorded. Moreover, even the type of S. caucasica fo.
desertorum (LE!) has slightly scabrous leaves. Thus, we
consider S. desertorum conspecific with S. glareosa.
During the revision of the specimens representing
Stipa glareosa we found interesting specimens collected
in southern Mongolia that represent this new variety,
which may also be found in the area of Middle Asia.
Stipa glareosa var. nemegetica M. Nobis, var. nov.
TYPE: southern Mongolia, southern Gobi Aimak,
Nemeget Mtns., alt. 2000–2700 m, 17 July 1964, K.
Kowalski s.n. (holotype, KRA 100264!).
Diagnosis. Stipa glareosa P. A. Smirn. var. nemegetica M.
Nobis differs from both varieties within S. glareosa (variety
glareosa and variety pubescens (P. A. Smirn. ex Roshev.)
Gubanov) in having scabrous vegetative and culm leaves
and very densely and long pubescent culm sheaths.
Stipa glareosa P. A. Smirn. var. pubescens (P. A. Smirn.
ex Roshev.) Gubanov, Byull. Moskovsk. Obshch.
Isp. Prir., Otd. Biol. 87(1): 124. 1982. Basionym:
Stipa glareosa fo. pubescens P. A. Smirn. ex
Roshev., Fl. URSS 2: 89. 1934. TYPE: Mongolia.
Gobi Altai, Bain-Tsagan, 4 Aug. 1931, N. P.
Ikonnikov-Galitzky & V. A. Ikonnikova-Galitzkaya
3824 (lectotype, designated by Grubov [1982], LE!).
Stipa glareosa var. pubescens, which is known only
from Mongolia, may also occur in Middle Asia. The
taxon differs from the nominal variety (S. glareosa var.
glareosa) in having leaves and sheaths densely and long
pubescent versus vegetative and culm leaves more or
less scabrous and sheaths of culm leaves that may be
glabrous or scabrous, respectively.
Stipa 3gnezdilloi Pazij, Opred. Rast. Sred. Azii 1:
201. 1968, pro sp. (S. caucasica Schmalh. 3S.
hohenackeriana Trin. & Rupr.). TYPE: Pamir-
Alai, Kh. Kugitang, zapadnyi sklon, okr. z.
Maidan, shchebnistye sklony, 20 July 1935, A.
Gnezdillo 134 (holotype, TASH 151761!; isotype,
TASH 151760!).
Habitat. Stipa 3gnezdilloi is found in steppe grass-
lands at 900–1400 m.
Distribution. Stipa 3gnezdilloi is found in Middle
Asia (Pazij, 1968), specifically in Uzbekistan (Kuhitang
Mountains). Figure 3.
Notes. The taxon is known only from the type col-
lection. In the original description, Pazij (1968) stated
that Stipa gnezdilloi has bigeniculate awns; however, the
holotype and isotype have unigeniculate or indistinctly
bigeniculate awns. The combination of morphological
characters (hairs on seta 3–4.2 mm, hairs on column
0.1–0.3 mm and gradually increasing toward genicula-
tion, as well as ligules of vegetative leaves that are ca.
0.4 mm long) suggests that the species originated from
hybridization between S. caucasica and S.hohenackeriana.
Stipa gracilis Roshev., Fl. Aziat. Ross. 1(12): 151, pl.
10, f. 4, 4a. 1916. TYPE: Semirechinskaya obl.
Pishpekckii uezd, dolina r. Chemaldynki, kame-
nistye sklony doliny, 23 July 1915, M. D. Spiri-
donow 17 (lectotype, designated by Nobis et al.
[2013: 670], MW!).
Habitat. Stipa gracilis is found in rock ledges and
fissures at 700–3000 m.
Distribution. Stipa gracilis is found in the moun-
tains of Middle Asia (Nobis & Nowak, 2011; Nobis
et al., 2013, 2014c). It occurs in southern Kazakhstan
(Tian-Shan Mountains), Kyrgyzstan (Alai Mountains,
Tian-Shan Mountains), northern Tajikistan (Pamir),
and western China (Tian-Shan Mountains). Figure 3.
Stipa grafiana Steven, Bull. Soc. Imp. Naturalistes
Moscou 30(2): 116. 1857. TYPE: [Ukraine.] Eka-
terinosevash Prov., 1851, Graff s.n. (holotype,
LE!). 5Stipa pulcherrima K. Koch subsp.
pulcherrima.
Stipa haussknechtii Boiss., Fl. Orient. [Boissier] 5(2):
501. 1884. TYPE: In rupibus calcareis montis
Sawers Persiae occid. supra Gulbar, 9000 ft., July
1868, Haussknecht s.n. (holotype, G not seen;
isotypes, JE!, W!, WU!). [Achnatherum hauss-
knechtii (Boiss.) M. Nobis, comb. nov.
Notes. Because of the lemma epidermis having a
maize-like pattern, dominated by silica bodies and short
fundamental cells (Fig. 7) typical for representatives of
Achnatherum rather than Stipa, the transfer of S. hauss-
knechtii to genus Achnatherum is needed. A similar
pattern in the lemma epidermis is observed in A.
mandavillei (Freitag) M. Nobis, comb. nov. (Fig. 7)
[Basionym: Stipa mandavillei Freitag, Davis & Hedge
Festschrift 118. 1989], which is known from Oman.
Stipa 3heptapotamica Golosk., Bot. Mater. Gerb.
Bot. Inst. Komarova Akad. Nauk S.S.S.R. 19:
46. 1959, pro sp. (S. lessingiana Trin. & Rupr.
28 Annals of the
Missouri Botanical Garden

3S. richteriana Kar. & Kir.; Nobis et al., 2019c).
TYPE: Yugo-zapadnye otrogi Dzhungarskogo Ala-
tau, gory Chulak, Monga-sai, po severnym oste-
pennym sklonam [SW spurs of Dzhungarian
Alatau, Chulak Mtns., Monga-sai, along south-
ern slopes], 29 May 1955, V. P. Goloskokov s.n.
[Typus!, I 1957, determ. V. P. Goloskokov] (lecto-
type, designated here, LE!; isolectotypes, LE!
[3 sheets]).
Habitat. Stipa 3heptapotamica is found in steppe
grasslands at 800–1500 m.
Distribution. Stipa 3heptapotamica is found in
southeastern Kazakhstan (Dzhungarski Alatau, north-
ern forelands of Tian-Shan Mountains; Goloskokov,
1959; Pazij, 1968; Tzvelev, 1976; Kotukhov, 2002;
Gudkova et al., 2015; Nobis et al., 2019c). Figure 4.
Notes. Stipa 3heptapotamica was described by
Goloskokov (1959) on the basis of one collection (south-
western spurs of Dzhungarskii Alatau, Chulak Mtns.,
Monga-sai, along southern slopes, 29 May 1955, V. P.
Goloskokov s.n.) composed of four herbarium sheets and
preserved at LE. In the description of the species,
Goloskokov (1959) included information that the type
is in LE; however, he did not select the holotype, but to
the specimens he attached additional labels: Typus I,
Typus II, and Typus III. The fourth specimen, which has
the same label as the three mentioned above, was
identified by Goloskokov as S. richteriana and redeter-
mined by Tzvelev in 1972 as another isotype of S.
heptapotamica. Tzvelev (1976) stated that the holotype
and three isotypes of this species are preserved in LE,
but he did not point out which one of these three
specimens is the holotype. Because a holotype of
S. 3heptapotamica was not formally chosen by the
author of the species, according to the International
Code of Nomenclature (McNeill et al., 2012; Turland
et al., 2018), a lectotype needs to be designated. Thus
the specimen with the above cited label and additional
labels “Typus! I, 1957, determ. V.P. Goloskokov”is
designated here as the lectotype.
Stipa himalaica Roshev., Bot. Mater. Gerb. Glavn. Bot.
Sada R.S.F.S.R. 5: 11. 1924. TYPE: [Pakistan.]
Gilgit Exped., S of Hindukush, Giles s.n. (lecto-
type, designated by Tzvelev [1968: 55], LE 9269!;
isolectotypes, E!, K!, WU!).
Habitat. Stipa himalaica is found in rocky ledges,
grasslands, and screes at 2000–4000 m.
Distribution. Stipa himalaica is found in the moun-
tains of central Asia (Pamir, Kunlun, Karakorum, Hin-
dukush, Himalayas; Freitag, 1985; Dickor ´e, 1995; Nobis
et al., 2013). In Middle Asia, it occurs in Afghanistan,
western China, and Pakistan (Fig. 4).
Stipa 3hissarica M. Nobis, Pl. Syst. Evol. 299(7):
1345. 2013 (S. lipskyi Roshev. 3S. orientalis
Trin.; Nobis, 2013). TYPE: Dolina rzeki Iskan-
derdaria (Iskanderdaya River valley), Rejon
Zerawsza ´nski B (Zeravshan Range B), Zachodni
Tad˙zykistan (western Tajikistan), Pamir Alai Mtns.,
Zeravshan Mtns., high mtn. steppe, among stones,
on the left slope of the Iskanderdarya River val-
ley, ca. 0.8 km ESE of Serimadarun Lake (near
Iskanderkul Lake), 39°059040N / 68°229520E,
elev. 2320 m, inclination SW, slope 5° to 10°, 14
June 2011, M. Nobis s.n. (holotype, KRA!; isotypes,
Herb. Stip. M. Nobis!, KRA).
Habitat. Stipa 3hissarica is found in high moun-
tain steppes at 2250–2350 m.
Distribution. Stipa 3hissarica is found in Middle
Asia (Nobis et al., 2013), specifically in Tajikistan
(Zeravshan Mountains).
Stipa hohenackeriana Trin. & Rupr., Sp. Gram.
Stipac. 80. 1842. TYPE: [Azerbaijan.] Transcau-
casia, circa Shusha et Helenendorf (Kirovabad),
Hohenacker 1253 (holotype, LE!).
Notes. Stipa hohenackeriana is a species highly var-
iable in morphology, and its taxonomic revision is needed.
Here, we recognize two varieties within the species.
1. Leaves of vegetative shoots scabrous, rarely almost
glabrous . . ................................
Stipa hohenackeriana Trin. & Rupr. var. hohenackeriana
19. Leaves of vegetative shoots pubescent . . .......
Stipa hohenackeriana var. grossheimii (Tzvelev) Tzvelev
Stipa hohenackeriana var. hohenackeriana.
Synonyms. 5Stipa pennata L. var. minor Boiss., Fl.
Orient. 5: 502. 1884, nom. illeg.; 5S. barbata Desf. var.
seminudata Hack., Denkschr. Kaiserl. Akad. Wiss.,
Wien. Math.-Naturwiss. Kl. 50: 8. 1885. TYPE: In
collibus ad viam versus Dauletabad (Malayer), Polak
882 (holotype, W!); 5S. atriseta Stapf ex Bor; 5S.
subbarbata B. Keller.
Habitat. Stipa hohenackeriana var. hohenackeriana
is found in sandy and stony steppes, grasslands, fallows,
and roadsides at 100–3000 m.
Distribution. Stipa hohenackeriana var. hohenack-
eriana is widely distributed from Asia Minor to central
Asia (Tzvelev, 1976, 2006; Freitag, 1985; Wu & Phil-
lips, 2006; Nobis et al., 2016b). In Middle Asia it
specifically occurs in Afghanistan, China, Iran,
Kazakhstan, Kyrgyzstan, Pakistan, Tajikistan, Turk-
menistan, and Uzbekistan.
Stipa hohenackeriana var. grossheimii (Tzvelev)
Tzvelev, Novosti Sist. Vyssh. Rast. 11: 16. 1974.
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Nobis et al. 29
Synopsis of Stipa (Poaceae) in Middle Asia

Basionym: Stipa hohenackeriana subsp. grossheimii
Tzvelev, Novosti Sist. Vyssh. Rast. 3: 21. 1966.
TYPE: Azerbaijan. Nakhichevanskaya ASSR, Nor-
ashenskii raion, otrogi g. Myunkh-Bala-Ogly, bliz
sel. Ulya-Norashen [Nakhichevan, Norashen Distr.,
Myunkh-Bala-Ogly Mtns.], ca. 900 m, 15 May
1947, A. A. Grossgeim, I. A. Ilinskaya & M. E.
Kirpichnikov s.n. (holotype, LE!; isotype, LE!).
Synonyms. 5Stipa hohenackeriana Trin. & Rupr.
subsp. ordubadica Tzvelev, Novosti Sist. Vyssh. Rast. 3: 22.
1966. TYPE: Ordubad region, kamenistyi sklon v nizhn.
gornom poyase Zangezyrskogo khr. k vost. ot Ordubada, 6
June 1956, T. Egorova, N. Tzvelev & S. Cherepanov s.n.
(holotype, LE!); [S. hohenackeriana var. ordubadica
(Tzvelev) Tzvelev, Novosti Sist. Vyssh. Rast. 11: 16. 1974.
Habitat. Stipa hohenackeriana var. grossheimii is
found in sandy and stony steppes at 400–2800 m.
Distribution. Stipa hohenackeriana var. grossheimii
is distributed within the range of S. hohenackeriana var.
hohenackeriana. In Middle Asia, it occurs in Afghani-
stan and Tajikistan (Pamir).
Stipa holosericea Trin., M´em. Acad. Imp. Sci. St.-
P´etersbourg, S ´er. 6, Sci. Math. 1: 81. 1830. TYPE:
[Iran, Azerbaijan.] In siccis montosis circa Bada-
lan, 8 June 1829, Szovits 410 (holotype, LE!).
Within the species, two varieties are recognized.
1. Adaxial surface of vegetative leaves covered with
mixture of short and long hairs, lower sheaths of
cauline leaves shortly pilose . . . . .............
.............. Stipa holosericea Trin. var. holosericea
19. Adaxial surface of vegetative leaves shortly pilose
with mixture of longer hairs on marginal ribs,
sheaths of lower cauline leaves glabrous . . . . . . .
Stipa holosericea var. transcaucasica (Grossh.) M. Nobis
Stipa holosericea var. holosericea.
Habitat. Stipa holosericea var. holosericea is found
in steppe grasslands at 1000–1500 m.
Distribution. Stipa holosericea var. holosericea is
found in southwestern Asia (Freitag, 1985). In Middle
Asia it occurs in northern Iran (Kopet-Dagh Mountains).
Stipa holosericea var. transcaucasica (Grossh.) M.
Nobis, comb. & stat. nov. Basionym: Stipa trans-
caucasica Grossh., Trudy Bot. Inst. (Baku) 2: 245.
1936.
Synonyms. [Stipa holosericea subsp. transcauca-
sica (Grossh.) Tzvelev, Novosti Sist. Vyssh. Rast. 11: 14.
1974.
Habitat. Stipa holosericea var. transcaucasica is
found in steppe grasslands at 800–1500 m.
Distribution. Stipa holosericea var. transcaucasica
is found in southwestern Asia (Tzvelev, 1976; Freitag,
1985; Nikitin & Geldikhanov, 1988). In Middle Asia, it
occurs in Turkmenistan (Kopet-Dagh Mountains).
Stipa ikonnikovii Tzvelev, Spisok Rast. Gerb. Fl. SSSR
21: 49. 1977. TYPE: Badachshan, ad ripam dex-
tram fl. Gunt, Czartym, in lapidosis, 5 Aug. 1957,
S. Ikonnikov s.n. [G. Ladygina, L. Sidorov] (holo-
type, LE!; isotypes, B!, JE!, KRAM!, MW!, TK!). [
Stipa kirghisorum P. A. Smirn. var. ikonnikovii
(Tzvelev) M. Nobis.
Stipa iljinii Roshev., Izv. Bot. Sada Akad. Nauk SSSR
30: 294. 1932. TYPE: Semipalatinskii u., okr. pos.
Znamenskogo, peschanaya kovylevaya step [Inter
Semipalatense, in steppis arenosis stipaceis non
procul a pago Snamenskoje], 6 July 1928, M. Ijlin
& O. Heinrichson 3867 (holotype, LE!; isotype,
LE!). [Stipa zalesskii Wilensky var. iljinii
(Roshev.) Tzvelev.
Stipa inebrians Hance, J. Bot. 14: 212. 1876. TYPE: Hab.
in montis Alashan, Mongoliae interioris, 1875,
Herb. propr., H. F. Hance 19204 (holotype, BM!;
isotype, LE!). [Achnatherum inebrians (Hance)
Keng ex Tzvelev, Rast. Tsentr. Azii 4: 40. 1968.
Stipa iraquensis Martinovsk´y, Preslia 42: 375. 1970.
TYPE: [Iraq.] 40 km ad orientem a Rutba positus,
27 Apr. 1961, Hadaˇc 4375 (holotype, PR!). 5
Stipa 3assyriaca Hand.-Mazz.
Stipa iskanderkulica (Tzvelev) Czerep., Sosud. Rast.
SSSR 387. 1981. Basionym: Stipa caucasica
Schmalh. subsp. iskanderkulica Tzvelev, Novosti
Sist. Vyssh. Rast. 11: 20. 1974. TYPE: Samar-
kand Distr., Iskanderkul, 21 May 1914, V.
Dubyanskii s.n. (holotype, LE!). [Stipa drobo-
vii (Tzvelev) Czerep. var. iskanderkulica (Tzve-
lev) M. Nobis & A. Nowak.
Stipa jacquemontii Jaub. & Spach., Ill. Pl. Orient. 4: 60, pl.
339. 1851. TYPE: [India.] Ad rupes in excelsis Emodi
Cashemyriani, 3000 m, Aug. 1831, V. Jacquemont 994
(lectotype, designated here, P00753746!; isolecto-
types, P00753747!, P00753745!, K000032095!). [
Achnatherum jacquemontii (Jaub. & Spach.) P. C.
Kuo & S. L. Lu, Fl. Reipubl. Popularis Sin. 9(3): 323.
1987; 5Achnatherum botschantzevii Tzvelev, Novosti
Sist.Vyssh.Rast.11:4.1974.TYPE:AsiaMedia,
jugum Alaicum, inter fl. Isfara et Soch, angustium
Kara-kulj austro-occidentaliter versus a pag. Kara-
bulak in rupibus calcareis, 15 July 1962, V. Botschant-
zev s.n. (holotype, LE!).
Notes. Because a holotype of Stipa jacquemontii was
not chosen formally by the authors of the species,
30 Annals of the
Missouri Botanical Garden

according to the International Code of Nomenclature
(McNeill et al., 2012; Turland et al., 2018), a lectotype
needs to be designated. Thus, the herbarium sheet with
specimens of S. jacquemontii, and the label mentioned
above including number P00753746, is designated here
as the lectotype.
Based on revised herbarium material of Achnatherum
botschantzevii collected from Kyrgyzstan and Uzbeki-
stan (northern part of the Alai Mountains) and preserved
at JE, KRA, and LE, we find this taxon conspecific with
previously described A. jacquemontii (Nobis et al.,
unpubl.). The locality of this species in the Alai Moun-
tains is the northernmost within the range of A.
jacquemontii.
Stipa jagnobica Ovcz. & Czukav., Izvest. Otdel.
Estestven. Nauk Akad. Nauk Tadzhik. SSR 17:
51. 1956 [1957]. TYPE: [Tajikistan.] Northern
slope of the Hissar Mtns., 5 km S of Takfon,
2600 m, 4 June 1949, Grigorev s.n. (holotype,
LE!; isotype, TAD!). [Stipa richteriana Kar.
& Kir. subsp. jagnobica (Ovcz. & Czukav.)
Tzvelev.
Stipa 3kamelinii Kotukhov, Turczaninowia 1(1): 10.
1998 (S. orientalis Trin. 3S. lessingiana Trin. &
Rupr.; Nobis & Gudkova, 2016). TYPE: Saur-
Tarbagataj, praemontium boreali-occidentale juge
Sajkan, in denudationibus tertiariis Akseir, jugi
parvi declivitas aqua erosa australi-occidentalis, in
glareoso-argillosis, 9 June 1992, Ju. Kotuchov s.n.
(holotype, LE!; isotypes, KRA 436045!, KRA
436046!, KUZ! [9 sheets]).
Habitat. Stipa 3kamelinii is found in steppe grass-
lands at 1200–1500 m.
Distribution. Stipa 3kamelinii is found in eastern
Kazakhstan (Tarbagatai Mountains; Kotukhov, 2002;
Nobis & Gudkova, 2016).
Notes. Morphologically, Stipa 3kamelinii is similar
to S. 3zaissanica Kotukhov, which grows in the southern
Altai Mountains; however, they differ in the length of the
ligules of the vegetative shoots, which are 0.3–1(–1.3) mm
versus (0.5–)3–6.5(–7.5) mm long, and the length of hairs on
the seta, which are 3.3–4 mm versus 2.3–3.6 mm, respec-
tively (Nobis & Gudkova, 2016). Kotukhov (1991, 1998a)
supposed that S. kamelinii arised as the result of the
hybridization of S. orientalis and S. 3zaissanica,whereas
S. zaissanica is a hybrid of S. orientalis and S. hohenack-
eriana. In our opinion it seems unlikely that S. 3kamelinii
originated from a backcrossing of S. 3zaissanica and S.
orientalis, since the ligules on the vegetative shoots of
S. 3kamelinii are much shorter than those in both
putative parental species (Nobis & Gudkova, 2016).
Stipa karakabinica Kotukhov, Bot. Zhurn. (Moscow &
Leningrad) 79(7): 105. 1994. TYPE: Altaj Australis,
jugum Tarbagatai, depressio Karakabinica, elev.
1800 m, moraenae clausae, declivitas substepposa
cum Sibiraea altaiensis, 18 Aug. 1986, Ju. Kotuchov
s.n. (holotype, LE!; isotypes, KRA 451780! [earlier
designated by Nobis & Gudkova, 2016: 34, as the
lectotype], KRA 436020!, KRA 436022!, KRA
436024!, KRA 436027!, KUZ! [4 sheets], LE! [2
sheets]).
Habitat. Stipa karakabinica is found in steppe
grasslands at 1700–1800 m.
Distribution. In Middle Asia Stipa karakabinica is
found in eastern Kazakhstan (Tarbagatai Mountains;
(Kotukhov, 2002; Nobis & Gudkova, 2016).
Notes. Although Kotukhov (1994) indicated the LE
herbarium as the place of the holotype, we were not able
to find it there during our research in 2008–2015 (Nobis
& Gudkova, 2016), thus the lectotype of the species was
designated from Kotukhov’s original collection, which is
preserved at KRA and KUZ. However, in December
2016 we found in LE three sheets with specimens of
S. karakabinica, with one of them being the holotype.
Stipa karakabinica is similar to S. capillata and
S. austroaltaica known from Altai Mountains (Nobis &
Gudkova, 2016), and probably both taxa mentioned
above are conspecific with S. capillata. Taxonomic revi-
sion of this group of taxa is required.
Stipa karataviensis Roshev., Trudy Pochv.-Bot.
Eksped. Izsl. Kolon. Raionov Aziatsk. Rossii,
Chast’II, Bot. Izsl. 6: 186. 1912. TYPE: Kazakh-
stan. Syr-Darin. obl.: Aulie-atinskii uezd 23 mai,
uschele Berk-kara (Karatau), no. 310, Iter ad distr.
Aulie-ata, 1909, Z. von Minkwitz s.n. (lectotype,
designated by Nobis [2013: 1329], LE!; isolecto-
types, LE!, W 35652!).
Habitat. Stipa karataviensis is found in steppe
grasslands at 700–1300 m.
Distribution. Stipa karataviensis is found in Middle
Asia (Pazij, 1968; Tzvelev, 1976; Nobis, 2010, 2013;
Gonzalo et al., 2012), specifically in southern Kazakh-
stan (Tian-Shan Mountains: Karatau, Dzhambyl, and
Chu-Illinskie Mountains) and Uzbekistan (Nuratau and
Aktau Mountains). Figure 4.
Stipa kazachstanica Kotukhov, Bot. Zhurn. (Moscow &
Leningrad) 79: 104. 1994. TYPE: Saur-Tarbagatai,
praemontia australi-occidentalia jugi Manrak, locus
Sarybulak, clivulus schistosus australi-orientalis,
12 June 1992, Ju. Kotuchov s.n. (holotype, LE!;
isotypes, LE!, KRA 436018!, KUZ! [2 sheets]). [
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Nobis et al. 31
Synopsis of Stipa (Poaceae) in Middle Asia

Stipa macroglossa P. A. Smirn. subsp. kazach-
stanica (Kotukhov) M. Nobis.
Stipa kempirica Kotukhov, Bot. Zhurn. (Moscow &
Leningrad) 79(7): 101. 1994. TYPE: Saur-Tarbagatai,
brachia australi-occidentalia jugi Manrak, elev. 600 m,
locus Kempirbulak, clivulus lapidosus orientalis, 11
July 1992, Ju. Kotuchov s.n. (lectotype, designated by
Nobis & Gudkova [2016: 34], LE!; isolectotypes,
KRA 436044!, KUZ!, LE!).
Habitat. Stipa kempirica is found in steppe grass-
lands at ca. 500–600 m.
Distribution. Stipa kempirica is found in eastern
Kazakhstan (Tarbagatai Mountains; Kotukhov, 2002;
Nobis & Gudkova, 2016).
Notes. This taxon is probably of hybrid origin; fur-
ther studies are required to identify its parental species.
Stipa kirghisorum P. A. Smirn., Repert. Spec.
Nov. Regni Veg. 21: 232–233. 1925. TYPE:
Semipalatinskaya obl., Bokai, Kossinskii s.n.
(holotype, MW!). EPITYPE: Semipalatinskaya
obl., Semipalatinskii uezd, Chingiz, zapadnaya
chast’gory Karagian-koi-tas, krutoi kamenistyi
sklon gory, 1 July 1914, C. Kossinsky 607
(epitype, designated by Nobis et al. [2016a:
151], LE!).
Within the species three varieties are recognized.
1. Awn glabrous or scabrous on column and plumose
on seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
19. Awn pilose throughout . . .....................
.....Stipa kirghisorum P. A. Smirn. var. balkhashensis
2. Ventral line of hairs on the anthecium terminating
0.3–1.0 mm below the top, adaxial surface of vege-
tative leaves with hairs 0.15–0.35 mm long . . . . . . .
.... Stipa kirghisorum var. ikonnikovii (Tzvelev) M. Nobis
29. Ventral line of hairs on the anthecium terminating
1–3(–4.6) mm below the top, adaxial surface of
vegetative leaves covered with prickles up to
0.1 mm long, sometimes with mixture of longer
hairs . . . . . . . . . . . . . Stipa kirghisorum var. kirghisorum
Stipa kirghisorum var. kirghisorum.
Synonyms. [Stipa pennata L. subsp. kirghisorum
(P. A. Smirn.) Freitag, Notes Roy. Bot. Gard. Edin-
burgh 42: 438. 1985; 5S. violacea Nikitina, hom.
illeg.; [S. nikitinae Tzvelev; [S. kirghisorum P. A.
Smirn. var. violacea Tzvelev, Novosti Sist. Vyssh. Rast.
11: 18. 1974.
Habitat. Stipa kirghisorum var. kirghisorum is
found in high mountain steppes, grasslands, screes,
and low scrubs at 350–3200 m.
Distribution. Stipa kirghisorum var. kirghisorum is
widely distributed in central Asia (Gonzalo et al., 2013;
Nobis et al., 2016a). In Middle Asia, it occurs in
Afghanistan, China, Kazakhstan, Kyrgyzstan, Pakistan,
Tajikistan, and Uzbekistan.
Stipa kirghisorum var. ikonnikovii (Tzvelev) M.
Nobis, comb. & stat. nov. Basionym: Stipa
ikonnikovii Tzvelev., Spis. Rast. Gerb. Fl.
SSSR 21(111–114): 49. 1977.
Habitat. Stipa kirghisorum var. ikonnikovii is found
in high mountain steppes and grasslands at 2000–2400 m.
Distribution. Stipa kirghisorum var. ikonnikovii is
found in Middle Asia, specifically in eastern Tajikistan
(Pamir Mountains; Tzvelev, 1977; Ikonnikov, 1979).
Stipa kirghisorum var. balkhashensis M. Nobis &
P. D. Gudkova, var. nov. TYPE: steppe grassland,
near the rd., near NW part of Balkhash Lake, 25 km
W of Balkhash City, 46°489040N / 74°419300E, alt.
360 m, wp. 251, 22 May 2014, M. Nobis &
P. Gudkova s.n. (holotype, KRA 432672!; isotype,
KRA 426156!). Figure 3.
Diagnosis. Stipa kirghisorum P. A. Smirn. var. balkha-
shensis M. Nobis & P. D. Gudkova differs from S. kirghisorum
var. kirghisorum only in having awns with shortly pubescent
(covered with 0.3–1 mm long hairs) versus glabrous and smooth
columns.
Habitat. Stipa kirghisorum var. balkhashensis is
found in steppe grasslands at ca. 350–400 m.
Distribution. Stipa kirghisorum var. balkhashensis
is found in Middle Asia, specifically in Kazakhstan
(near northwest part of Balkhash Lake).
Notes. See comments under Stipa borysthenica var.
anomala.
Stipa koenigii Woronow, Bot. Mater. Gerb. Glavn. Bot.
Sada R.S.F.S.R. 5: 61. 1924. TYPE: Culta in
sectione caucasica Horti Tiflisiensis e seminbus a
cl. E. Koenig e distr. Olty prov. Kars a. 1906 allatis,
Yu. Woronow s.n. (holotype, LE!). 5Stipa arabica
Trin. & Rupr. var. turgaica (Roshev.) Tzvelev.
Stipa 3kolakovskyi Tzvelev, Bot. Zhurn. (Moscow &
Leningrad) 78(10): 93. 1993. TYPE: Norashenskii
r-n, na kholmakh okolo sel. Diza, na sev. sklone, 4
June 1947, A. Grossheim, I. Ilinskaya, M. Kirpich-
nikov & Gerb. Fl. SSSR 3666 (holotype, LE!). 5
Stipa 3assyriaca Hand.-Mazz.
Stipa kopetdaghensis Czopanov, Novosti Sist. Vyssh.
Rast. 6: 22. 1970. TYPE: Central Kopetdag Mtns.,
western slope, near the top of Dushak Mtn., 30 July
1967, P. Czopanov s.n. (holotype, ASH not seen;
isotype, LE!). 5Stipa 3alaica Pazij.
32 Annals of the
Missouri Botanical Garden

Stipa korshinskyi Roshev., Fl. Aziat. Ross. 1(12): 163, t.
11. 5, 5a. 1916. TYPE: Kazakhstan. Akmolinskaya
obl., Atbasarskii u., tipchakovo-kovyl’naya step v okr.
st. Atbasarskoi, 30 May 1908, no. 2, V. Kapelkin s.n.
(lectotype, designated by Tzvelev [1976: 578], LE!).
Habitat. Stipa korshinskyi is found in steppes and
grasslands at 300–1000 m.
Distribution. Stipa korshinskyi is found in Kazakh-
stan (Aral-Caspian and Balkhash regions) and southern
Russia (southern Ural, Transvolga, western Siberia; Tzve-
lev, 1976; Punina & Gudkova, 2016). In Middle Asia, it
occurs in central Kazakhstan (around Balkhash Lake).
Notes. Because of its shortly pilose awns, with hairs
on the seta 0.3–0.5 mm long, Stipa korshinskyi is similar
to S. richteriana s.l. However, the latter taxon differs
from S. korshinskyi by having straight setae, up to 5 cm
long versus setae more or less flexuous and longer than
5 cm; callus (0.5–)0.6–1.1 mm long and bent versus
callus 1.3–1.8 mm long and straight. The possibility
should not be excluded that S. korshinskyi originated
from hybridization between S. sareptana and S. rich-
teriana subsp. richteriana. Further studies are needed
to evidence this; however, during field studies in
Kazakhstan in 2019, we found two small populations
of S. korshinskyi growing within populations of the two
species mentioned above.
New records. KAZAKHSTAN. Northern part of Dzhungar-
skii Alatau, N Tian-Shan, ca. 10 km NEE of Saryozek, NE of
Almaty, steppe, 44°239060N / 78°059520E, 936 m, wp. 1254,
27 May 2019, M. Nobis s.n. (KRA); 63 km NW of Usharal, E of
Balkhash Lake, steppe, 46°319320N / 80°179290E, 404 m, wp.
1226, 23 May 2019, M. Nobis s.n. (KRA).
Stipa kotuchovii M. Nobis, Ann. Bot. Fenn. 48: 494.
2011 [replaced synonym: Stipa monticola Kotu-
khov, hom. illeg.]. TYPE: Saur-Tarbagataj, brachia
austro-occidentalia jugi Saur, in viciniis hiberna-
culi Kyzylkija, elev. 1700 m, declivitas australi-
orientalis, 18 Aug. 1992, Ju. Kotuchov s.n. (holo-
type, LE!; isotypes, KRA 436032!, KRA 435918!,
KRA 436039!, KRA 436040!, KUZ! [2 sheets]).
Habitat. Stipa kotuchovii is found in steppe grass-
lands at 1600–1700 m.
Distribution. Stipa kotuchovii is found in Middle
Asia (Kotukhov, 2002; Nobis & Gudkova, 2016), spe-
cifically in eastern Kazakhstan (Tarbagatai Mountains).
Notes. Stipa kotuchovii is very similar to S. sczerba-
kovii Kotukhov, and most probably it should be treated
as conspecific with the latter taxon. Kotukhov (1998a)
distinguished it from S. sczerbakovii by its longer anthe-
cium (9.5–11.5 vs. 8.5–9 mm long), shorter awn (7–9 vs.
9–10 mm long), and vegetative leaves (adaxial surface
covered by mixture of short and long hairs vs. adaxial
surface covered only by short hairs). However, in ac-
cordance with the results of our revision, variability of
particular characters of S. sczerbakovii is much greater
than was detailed in the protologue (see comment under
S. sczerbakovii). Because both of these taxa probably
have a hybrid origin, taxonomical revision using mo-
lecular methods is needed.
Stipa krascheninnikowii Roshev., Mat. Komiss. Eksped.
Issl. Akad. Nauk, Ser. Kazakst. 5: 253. 1928.
TYPE: Aktyubinskaya obl., bass. Khobdy, sklon
k Tamde, kovylno-raznotravnaya step, 7 July 1926,
I. Ilin & I. Avramchik 297 (lectotype, designated by
Tzvelev [1976: 588], LE!). 5Stipa ucrainica
P. A. Smirn.
Stipa krylovii Roshev., Izv. Glavn. Bot. Sada S.S.S.R.
28: 379. 1929. TYPE: Selenginskaya Dauriya, gory
mezhdu Temnikom i Dzhidoi, yugo-zapadnaya
chast khr. Borgoiskogo, na sklonakh so stepnoi
rastitelnostyu, 28 July 1912, V. Smirnov 524 (lec-
totype, designated by Tzvelev [1976: 581], LE!).
Synonyms. [Stipa sareptana A. K. Becker subsp.
krylovii (Roshev.) D. F. Cui, Fl. Xinjiangensis 6: 299.
1996; [S. sareptana var. krylovii (Roshev.) P. C. Kuo &
Y. H. Sun, Fl. Reipubl. Popularis Sin. 9(3): 275. 1987;
5S. capillata L. var. coronata Roshev., Fl. Aziat. Ross.
12: 168. 1916. TYPE: LE!; 5S. densiflora P. A. Smirn.;
5S. densa P. A. Smirn.; 5S. decipiens P. A. Smirn.
Habitat. Stipa krylovii is found in steppes, screes,
and sandy and rocky grasslands at 500–3500 m.
Distribution. Stipa krylovii is widely distributed in
central Asia. Its occurrence is known in Russia, Mon-
golia, China, Kazakhstan, Kyrgyzstan, Tajikistan, India,
and Nepal (Pazij, 1968; Tzvelev, 1968, 1976; Freitag,
1985; Wu & Phillips, 2006; Nobis et al., 2016c;
Gudkova et al., 2017a, 2017b). In Middle Asia it occurs
in China, eastern Kazakhstan (Tarbagatai Mountains,
Tian-Shan Mountains), Kyrgyzstan (central Tian-Shan
Mountains), and Tajikistan (Pamir). Figure 8.
Stipa kuhitangi Drobow, Fl. Uzbekist. 1: 537. 1941.
TYPE: Greben Kugitanga protiv s. Kyzyl-alma, 27
June 1927, M. G. Popov 157 (holotype, TASH
47030!). 5Stipa richteriana Kar. & Kir. var.
richteriana.
Stipa kungeica Golosk., Bot. Mater. Gerb. Bot. Inst.
Komarova Akad. Nauk S.S.S.R. 16: 39. 1954. [
Stipa macroglossa P. A. Smirn. var. kungeica
(Golosk.) M. Nobis.
Stipa kyzylkiensis Kotukhov, Turczaninowia 1(2): 12–
13. 1998. TYPE: Saur-Tarbagataj, jugum Saur, in
regione hibernacula Kyzylkija, declivitas australi-
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Nobis et al. 33
Synopsis of Stipa (Poaceae) in Middle Asia

orientalis lapidosa, 17 July 1993, Ju. Kotuchov s.n.
(lectotype, designated by Nobis & Gudkova [2016:
35], LE!; isolectotypes, KRA 436041!, KRA
436042!, KUZ! [6 sheets], LE! [2 sheets]). 5Stipa
sczerbakovii Kotukhov.
Stipa langshanica (Y. Z. Zhao) Y. Z. Zhao, Acta Sci.
Nat. Univ. Neimenggu. 27(2): 211. 1996. TYPE:
China. Nei Monggol, Bayannaoermen, Langshan,
in clivis, 4 July 1988, Zhao et al. 4077 (holotype,
HIMC not seen). 5Stipa glareosa P. A. Smirn.
Stipa laxiflora Keng, Sunyatsenia 6(1): 73. 1941.
TYPE: [China.] Kansu [Gansu] Prov., 17 Oct.
1934, C. W. Yao 562 (Herbarium of the Biological
Laboratory, the Science Society of China no.
49597, not seen). 5Stipa penicillata Hand.-
Mazz. [syn. after Tzvelev, 1968].
Stipa lessingiana Trin. & Rupr., Sp. Gram. Stipac. 79.
1842. TYPE: “Stipa pennata L.,”Orenburg
Gubern., 1834, Lessing 413 (holotype, LE Herbar-
ium Trinii 1413.1!).
1. Sheaths of cauline leaves glabrous . . . . . . . . . . . . .
...... Stipa lessingiana Trin. & Rupr. var. lessingiana
19. Sheaths of cauline leaves pubescent . . . . . . . . . . .
........ Stipa lessingiana var. brauneri (Pacz.) Roshev.
Stipa lessingiana var. lessingiana.
Synonyms. 5Stipa saikanica Kotukhov.
Habitat. Stipa lessingiana var. lessingiana is found
in mountain steppes, grasslands, rocks, and gravelly
slopes at 300–1500 m.
Distribution. Stipa lessingiana var. lessingiana is a
widely distributed species. Its range extends from eastern
Europe (Greece, Bulgaria, Romania, Ukraine) through
Turkey, Georgia, Armenia, Azerbaijan, Iran, Kazakhstan,
and southern Russia up to western China (Pazij,
1968; Tzvelev, 1968, 1976; Klokov & Osychnyuk,
1976; Martinovsk´y, 1980; Freitag, 1985; Nobis et al.,
2019a). In Midd le Asia, it occur s in Ch ina, Ir an,
Kazakhstan, Kyrgyzstan, Tajikistan, Turkmenistan, and
Uzbekistan. Figure 9.
Stipa lessingiana var. brauneri (Pacz.) Roshev., Fl.
URSS 2: 93. 1934. Basionym: Stipa lessingiana
Trin. & Rupr. subsp. brauneri Pacz., Zap. Krymsk.
Obsch. Estestvoips. 5: 4. 1916. TYPE: Tauria,
penins, Tarchankut, prope Ak-Meczet, 30 Apr.
1914, A. Brauner s.n. (holotype, LE!).
Synonyms. [Stipa brauneri (Pacz.) Klokov.
Habitat. Stipa lessingiana var. brauneri is found in
steppes and grasslands at 300–1000 m.
Distribution. Stipa lessingiana var. brauneri was
noted within the range of the nominal variety (Ukraine,
Kazakhstan, southwestern Russia, Georgia; Tzvelev,
1976, 2006; Klokov & Osychnyuk, 1976). In Middle
Asia it occurs in Kazakhstan.
Figure 8. Distribution maps. Stipa krylovii Roshev. (=) and S. sareptana A. K. Becker (¤).
34 Annals of the
Missouri Botanical Garden

Stipa lingua A. Junge, Bull. Jard. Bot. Petersb. 10:
129. 1910. TYPE: [Turkmenistan.] Flora turko-
manica, prope Germab, 30 May 1889, A. Antonov
(holotype, LE!; isotype, LE!, MW!).
Synonyms. 5Stipa lingua A. Junge fo. brevifolia
Roshev., Fl. Aziat. Ross., 12: 146. 1916. TYPE: Tajiki-
stan. Schugnan, ˇ
Sachdary Valley, betw. Drum Dar River
& Chay-Kil River, 26 July 1914, Tuturin & Bessedin
371 (lectotype, designated by Gonzalo et al. [2011:
415], LE!); 5S. platypoda Bor.
Habitat. Stipa lingua is found in mountain steppes,
grasslands, deserts, screes, and gravelly slopes at
1000–2600 m.
Distribution. Stipa lingua is found in the southern
part of Middle Asia (Freitag, 1985; Gonzalo et al.,
2012). It also occurs in Afghanistan, Iran, Tajikistan,
Turkmenistan, and Uzbekistan. Figure 3.
Stipa lipskyi Roshev., Fl. Aziat. Ross. 12:153, pl. 10, f.
5, 5a. 1916.TYPE: [Middle Asia.] SrednyayaAziya,
Samarkand obl., Samarkand, 27 May 1897, V. I.
Lipskii 4530 (lectotype, designated by Tzvelev
[1976: 594], LE!; isolectotypes, K!, MW!).
Within the species two varieties are recognized.
1. Upper cauline sheaths glabrous to slightly sca-
brous . . . . . . . . . . . . . . Stipa lipskyi Roshev. var. lipskyi
19. Upper cauline sheaths pilose along the veins . . .
............ Stipa lipskyi var. pilosivaginata M. Nobis
Stipa lipskyi var. lipskyi.
Synonyms. [Stipa lingua A. Junge subsp. lipskyi
(Roshev.) R. Gonzalo, Ann. Bot. Fenn. 48(2): 159. 2011.
Habitat. Stipa lipskyi var. lipskyi occurs in moun-
tain steppes, grasslands, deserts, and gravelly slopes at
300–2400 m.
Distribution. Stipa lipskyi var. lipskyi is found in
Middle Asia (Nobis, 2013) in Kazakhstan, Kyrgyzstan,
Tajikistan, and Uzbekistan.
Stipa lipskyi var. pilosivaginata M. Nobis, Pl. Syst.
Evol. 299(7): 1338. 2013. TYPE: Tajikistan. Zer-
avshan Mtns., high mtn. steppe, on the left slope of
Fan River valley, near the N part of Zeravshan II
settlement, 39°119460N / 68°329040E, alt. 1750 m,
exp. NE, slope 30° to 50°, no. 5, 16 June 2012, M.
Nobis s.n. (holotype, KRA 383077!; isotypes, Herb.
Stip. M. Nobis!, KRA!, KRAM!).
Habitat. Stipa lipskyi var. pilosivaginata is found in
arid, high mountain steppes at 1700–1800 m.
Distribution. Stipa lipskyi var. pilosivaginata is
found in Tajikistan (Zeravshan Mountains; Nobis, 2013).
Stipa litwinowiana P. A. Smirn. ex Pavlov & Lipsch., Sovietsk.
Bot. 19. 1934. Replaced synonym: Oryzopsis turcoman-
ica Roshev., Fl. Aziat. Ross. 12: 184. 1916. TYPE:
Turcomania, in montibus supra Firuza prope Chan-
Jaila, 1200 m, 18 July 1897, D. Litvinov 184a (holotype,
Figure 9. Distribution maps. Stipa lessingiana Trin. & Rupr. (¤), S. margelanica P. A. Smirn. (s), and S. 3pseudoma-
croglossa M. Nobis (□).
Volume 105, Number 1
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Nobis et al. 35
Synopsis of Stipa (Poaceae) in Middle Asia

LE!). [Achnatherum turcomanicum (Roshev.)
Tzvelev, Novosti Sist. Vyssh. Rast. 11: 6. 1974.
Stipa longiplumosa Roshev. ex Kom., Fl. URSS 2:
87–88. 1934. TYPE: Khrebet Sarsaryak,vostochnyi
sklon okrestnosti kishl. Margak, kholmy nizhe Mar-
gaka, v 2–3 km ot nego na Yu-V, no. 227, 7 June
1932, N. Gontscharow, G. Grigorjev & V. Nikitin s.n.
(lectotype, designated by Nobis [2013: 1332], LE!;
isolectotypes, LE!).
Habitat. Stipa longiplumosa is found in mountain
steppes, grasslands, deserts, and gravelly slopes at
900–1500 m.
Distribution. Stipa longiplumosa is found in Middle
Asia (Nobis, 2013), specifically in Kyrgyzstan, Tajiki-
stan, and Uzbekistan (Fig. 4).
Notes. Stipa longiplumosa, with its description in
Russian, was published by Roshevitz (1934) who refer-
enced its Latin description to another publication (Acta
Inst. Bot. Acad. Sci. URSS 2[2]. 1934). However, the Latin
description of the species was not actually published until
two years later in 1936 (Roshevitz, 1936; Nobis, 2013).
Stipa macroglossa P. A. Smirn., Bot. Mater. Gerb. Glavn.
Bot. Sada R.S.F.S.R. 5: 47. 1924. TYPE: I. M.
Krasheninnikov [sic]: Ekspeditsiya v Turgaiskii uezd
1914 g., no. 5203, Turgaiskaya obl. i u. Kizil-
dzhingilskaya volost, r. Sary-su v svoikh nizovyakh,
Okrestnosti Muyun-kumov, Obnazhenie tretichnykh
peschanikov, 1 June 1914, H. Krascheninnikov
s.n. [sic] (lectotype, designated by M. Nobis in
Nobis et al. [2014a: 84], LE!; isolectotypes, LE!
[3 sheets]).
Stipa macroglossa subsp. macroglossa.
Synonyms. [Stipa turkestanica Hack. subsp. mac-
roglossa (P. A. Smirn.) R. Gonzalo, Syst. Bot. 38: 370.
2013.
Within the taxon two varieties are recognized.
1. Sheaths of cauline leaves glabrous . . . . . . . . . . . . .
........ Stipa macroglossa P. A. Smirn. var. macroglossa
19. Sheaths of cauline leaves pubescent . . . . . . . . . . .
Stipa macroglossa var. pubescens (P.A.Smirn.)M.Nobis
Stipa macroglossa P. A. Smirn. var. macroglossa.
Habitat. Stipa macroglossa var. macroglossa is
found in mountain steppes, grasslands, and screes at
1100–2400 m.
Distribution. Stipa macroglossa var. macroglossa
is found in Middle Asia (Gonzalo et al., 2013;
Nobis et al., 2016a), specifically southern Kazakh-
stan, Kyrgyzstan, Tajikistan, and Uzbekistan.
Stipa macroglossa var. pubescens (P. A. Smirn.) M.
Nobis, Pl. Syst. Evol. 302: 151. 2016a. Basionym:
Stipa macroglossa fo. pubescens P. A. Smirn., Bot.
Mater. Gerb. Glavn. Bot. Sada R.S.F.S.R. 5: 48.
1924. TYPE: Semirech. obl., Przhev. u. Ur. Toguz-
torau, terrasy prav. stor. r. Kugart, 9 June 1913,
V. Sapozhnikov 35 (lectotype, designated by
Nobisetal. [2016a: 151], LE!; isolectotype, LE!).
Habitat. Stipa macroglossa var. pubescens is found
in mountain steppes at 1500–2000 m.
Distribution. Stipa macroglossa var. pubescens is
found in Middle Asia (Nobis et al., 2016a) in southern
Kazakhstan and Tajikistan.
Stipa macroglossa subsp. kazachstanica (Kotukhov)
M. Nobis, Pl. Syst. Evol. 299: 1352. 2013. Basio-
nym: Stipa kazachstanica Kotukhov, Bot. Zhurn.
(Moscow & Leningrad) 79: 104. 1994.
Within the taxon two varieties are recognized.
1. Awn glabrous or scabrous on column and plumose
on seta . . . . . . . . . . . . . . . . . . . . Stipa macroglossa
P. A. Smirn. var. kazachstanica (Kotukhov) M. Nobis
19. Awn pilose throughout . . . . . ..................
.... Stipa macroglossa var. kungeica (Golosk.) M. Nobis
Stipa macroglossa var. kazachstanica (Kotukhov) M.
Nobis, comb. & stat. nov. Basionym: Stipa kazach-
stanica Kotukhov, Bot. Zhurn. (Moscow & Leningrad)
79: 104. 1994.
Habitat. Stipa macroglossa var. kazachstanica is
found in mountain steppes, grasslands, and screes at
1000–2800 m.
Distribution. Stipa macroglossa var. kazachstanica
is found in the mountains of central Asia (Kotukhov,
2002; Nobis et al., 2016a, 2016b). In Middle Asia, it
occurs in western China, eastern Kazakhstan, and
Kyrgyzstan. Figure 3.
Stipa macroglossa var. kungeica (Golosk.) M. Nobis,
comb. & stat. nov. Basionym: Stipa kungeica
Golosk., Bot. Mater. Gerb. Bot. Inst. Komarova
Akad. Nauk. S.S.S.R. 16: 39. 1954. TYPE: [Kazakh-
stan.] Severnye otrogi Kungei-Alatau, Tau-Chilik, v
1 km nizhe vpadeniya r. Kaindy, po yuzhnomu kame-
nistomu sklonu pestrotsvetnykh tolysch [northern
spurs of Kungei Alatau, Tau-Chilik, 1 km below
the fall of Kainda, along southern stony slopes], 9
June 1953, V. P. Goloskokov s.n. [“TYPUS I, V.
Goloskokov”and “Stipa kingeica m. Typus!, II
[Feb.] 1954, determ. V. P. Goloskokov”] (lecto-
type, designated here, LE!; isolectotypes, AA!,
LE! [2 sheets]).
36 Annals of the
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Habitat. Stipa macroglossa var. kungeica is found in
steppes and grasslands at 1000–1500 m.
Distribution. Stipa macroglossa var. kungeica is
found in Middle Asia (Goloskokov, 1954; Pazij,
1968; Tzvelev, 1976), specifically in Kazakhstan
(Dzhungarski Alatau).
Notes. Stipa macroglossa var. kungeica is known
only from the type collection. Morphologically it is
close to S. macroglossa var. kazachstanica, but dif-
fers in having pubescent (vs. glabrous) awn columns.
Stipa kungeica was described by Goloskokov (1954) on
the basis of one collection (1 km below the fall of Kainda,
along southern stony slopes, 9 June 1953, V. P.
Goloskokov s.n.) that is preserved on four sheets, three
at LE and one at AA. In the description of the species,
Goloskokov (1954) included information that the type is
preserved at LE. However, he did not indicate the
holotype, but to the specimens he attached additional
labels: Typus I, Typus II, and Typus III. Tzvelev
(1976) stated that the holotype and two isotypes of
this species are preserved at LE, but he did not point
out which one of these three specimens was the ho-
lotype. Because the holotype of S. kungeica was not
formally chosen by the author of the species, according
to the International Code of Nomenclature (McNeill
et al., 2012; Turland et al., 2018), the lectotype needs
to be designated. Thus the specimen with the above
cited label and two additional labels, “TYPUS I, V.
Goloskokov.”and “Stipa kingeica m. Typus!, II 1954,
determ. V. P. Goloskokov,”is designated here as the
lectotype.
Stipa maeotica Klokov & V. V. Osychnyuk, Novosti Sist.
Vyssh. Nizsh. Rast. 1975: 60–61. 1976. TYPE:
[Ukraine.] Donetskaya obl., Novoazovskii r-n,
Khomutovskaya step, 11 June 1955, F. Grin s.n.
(holotype, KW; isotype, LE!). 5Stipa zalesskii
Wilensky var. zalesskii.
Stipa magnifica A. Junge, Bull. Jard. Bot. Petersb. 10:
128. 1910. TYPE: Turkestania, Prov. Fergana,
Distr. Osch, prope Gulcza, 7 June 1900, W. Trans-
chel s.n. (holotype, LE!; isotypes, LE!, MW!).
Synonyms. [Stipa lingua A. Junge subsp. magnifica
(A. Junge) R. Gonzalo, Ann. Bot. Fenn. 48(2): 160.
2011; [S. caucasica Schmalh. var. tranzschelii
Drobow, Repert. Spec. Nov. Regni Veg. 21: 37.
1925. TYPE: probably the same as for Stipa mag-
nifica;5S. barbata Desf. var. platyphylla Hack.,
Vidensk. Meddel. Dansk Naturhist. Foren. Kjøben-
havn 55: 163. 1903. TYPE: Kyrgyzstan. Alai Mtns.,
Sufi Kurgan, 18 July 1898, Paulsen 407 (holotype, C
not seen, syn. after Gonzalo et al. 2011).
Habitat. Stipa magnifica is found in mountain
steppes, grasslands, and screes at 600–1900 m.
Distribution. Stipa magnifica occurs in Middle Asia
(Tzvelev, 1976; Gonzalo et al., 2012; Nobis, 2012;
Lazkov & Sultanova, 2014), specifically in southwestern
Kyrgyzstan (Fig. 3).
Stipa 3manrakica Kotukhov, Bot. Zhurn. (Moscow &
Leningrad) 74(3): 414. 1989 (Stipa caucasica
Schmalh. 3Stipa macroglossa P. A. Smirn. subsp.
kazachstanica (Kotukhov) M. Nobis; Nobis, 2010).
TYPE: Kazachstania Orientalis, jugi Manrak pars
centralis, locus Sagyndyk Major, in regione
montana media, 1100 m supra mare, jugi parvi
declivitas occidentalis schistosa, 27 June 1998,
J. Kotuchov s.n. (lectotype, designated by Nobis
[2010: 737], LE!; isolectotypes, KUZ! [2 sheets],
LE! [2 sheets]).
Synonyms. 5Stipa saurica Kotukhov.
Habitat. Stipa 3manrakica is found in steppe
grasslands at 1500–2600 m.
Distribution. Stipa 3manrakica occurs in Middle
Asia (Kotukhov, 2002; Nobis, 2010, 2012; Nobis et al.,
2015a), specifically in Kazakhstan (Tarbagatai
Mountains, Tian Shan Mountains) and Kyrgyzstan
(Tian-Shan Mountains).
Stipa margelanica P. A. Smirn., Repert. Spec. Nov.
Regni Veg. 26: 264. 1929. TYPE: [Kyrgyzstan.]
Ferg. obl. Margel. u., mezhdu dol. r. Isfairam i ur.
Khodzha-aryk, sukhoi kamenistyi zap. sklon, u
perevala [Fergana, Distr. Margelan, inter val. fl.
Isfairam et pread. Chodsha-aryk, ad trajectum in
declivitate sicca petrosa], 31 May 1913, N. Des-
siatoff 892 (lectotype, designated here, LE!; iso-
lectotypes, LE!, MW!).
Habitat. Stipa margelanica isfoundinsteppe
grasslands at 1000–2000 m.
Distribution. Stipa margelanica occurs in Middle
Asia (Ovchinnikov & Chukavina, 1957; Pazij, 1968;
Tzvelev, 1976; Freitag, 1985), specifically in southern
Kyrgyzstan, Tajikistan, and Afghanistan.
Notes. The original material representing Stipa
margelanica (type) was prepared on the basis of one
collection, no. 892, leg. N.Dessiatoff, which is pre-
served on two herbarium sheets at LE and one at MW.
On each of the herbarium sheets deposited at LE,
Smirnov attached his revision label “Stipa macroglossa
P. Smirn., 1927.VI, Det. P.A. Smirnow.”In addition, on
both sheets white labels with the inscription Typus have
been attached by herbarium employees, but only one of
them has an additional label in the left corner: “Kew
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Nobis et al. 37
Synopsis of Stipa (Poaceae) in Middle Asia

negative, no. 6921, 4 Mar 1965.”It is likely that based
on this, Tzvelev (1976) treated that sheet as the type (5
holotype), regarding the second one as an isotype.
Because the holotype was not chosen by the author of
the species description, the lectotype, according to the
International Code of Nomenclature (McNeill et al.,
2012; Turland et al., 2018), still needs to be designated.
Thus, the specimen on the sheet with the label “Kew neg-
ative...”is designated here as a lectotype of the species. The
other specimens cited from LE and MW are isolectotypes.
Stipa minuscula F. M. V´azquez, Telopea 13(1–2): 166,
f. 1k, m. 2011. TYPE: Tibet. Plants of the NE part
of the Qinghai-Xizang (Tsinghai-Tibet) Plateau,
C.G. 81-0362 8, s. coll. (holotype, BM001191177!
[single specimen in the upper left corner of the
sheet]). 5Stipa glareosa P. A. Smirn. var. glareosa.
Stipa mongholica Turcz. ex Trin., M´em. Acad. Imp. Sci.
Saint-P´etersbourg, S´er. 6, Sci. Math., Seconde Pt.
Sci. Nat. 4: 42. 1836. TYPE: Stipa mongholica
Turcz, Transbaical (coll. of 12 syntypes at LE!, cf.
Sokolova, 2012). [Ptilagrostis mongholica
(Turcz. ex Trin.) Griseb., Fl. Ross. 4(13): 447.
1852 [[Achnatherum mongholicum (Turcz. ex
Trin.) Ohwi, J. Jap. Bot. 17: 403. 1941].
Notes. The data on the distribution of Ptilagrostis
mongholica in Kyrgyzstan and Tajikistan (given, e.g., by
Chukavina & Ovchinnikov, 1957; Lazkov & Sultanova,
2014) should be referred to P. malyschevii Tzvelev
(Novosti Sist. Vyssh. Rast. 11: 7–8. 1974. TYPE:
Tjanj-Schanj centralis, in declivitatibus borealibus ad
fl. Buzulgan, 18 July 1908, R. Rozhevitz 1244 [holotype,
LE!; isotype, LE!]), whose occurrence in this region was
confirmed by us during both field studies and revision of
the herbarium materials.
Stipa monticola Kotukhov, Turczaninowia 1(1): 11.
1998, hom. illeg. non Stipa monticola H. Scholz,
Willdenowia 23(1–2): 117. 1993. TYPE: Saur-
Tarbagataj, brachia austro-occidentalia jugi Saur,
in viciniis hibernaculi Kyzylkija, elev. 1700 m,
declivitas australi-orientalis, 18 Aug. 1992, Ju.
Kotuchov s.n. (holotype, LE!; isotypes, KRA
436032!, KRA 435918!, KRA 436039!, KRA
436040!, KUZ! [2 sheets]). [Stipa kotuchovii
M. Nobis.
Stipa nakaii Honda, Rep. First Sci. Exped. Manchou-
kuo 4(4): 104. 1936. TYPE: Manshuria, Prov.
Chin-chou, Prope Chao-yang, 8 Aug. 1933, Nakai,
Honda & Kitagawa s.n. (holotype, KYO not seen).
[Achnatherum nakaii (Honda) Tateoka ex
Imzab, Fl. Intramong. 7: 196. 1983.
Stipa narynica M. Nobis, Nordic J. Bot. 30(1): 70–72,
f. 1, 3 1a–d, 4a. 2012. TYPE: Western Kyrgyzstan,
western Tian-Shan Mtns., Naryn River valley,
steppe grasslands on the right calcareous slope
of stream valley, a left tributary of the Naryn River,
in the eastern part of Tash Kumyr town, 41°199N/
72°129E, alt. 650 m, exposition S, slope 40°, no. 1,
11 May 2011, M. Nobis & A. Nowak s.n. (holotype,
KRA 382227!; isotypes, FRU!, Herb. Stip. M.
Nobis!, KRA!, KRAM!).
Habitat. Stipa narynica is found in steppe grass-
lands at 500–1200 m.
Distribution. Stipa narynica occurs in Middle Asia
(Nobis, 2012), specifically in western Kyrgyzstan (Chat-
kal and Fergana Mountains). Figure 4.
Stipa nikitinae Tzvelev, Novosti Sist. Vyssh. Rast. 43:
27. 2012. TYPE: Tsentralnyi Tian-shan, severnyi
sklon khr. Kavak-tau, pereval mezhdu Beiryuk i
Kash-bel, 28 July 1937, E. Mikhailova & L. Popova 66
(holotype, LE!). 5Stipa kirghisorum P. A. Smirn.
Stipa okmirii Dengub., Bot. Zhurn. (Moscow & Leningrad)
65(3): 431. 198 0. TYPE: Jugum Schugnanicum, in
viciniis horti botanici Pamirensis, declivia lapi-
dosa ad austro-occidentem exposita [Tadzhikistan,
GBAO, Shugnanskii khr., okrestnosti botaniche-
skogo sada, sklon], elev. 2350 m, 14 June 1977, A.
V. Dengubenko 2731 (holotype, LE!).
Habitat. Stipa okmirii is found in steppe grasslands
at 2250–2500 m.
Distribution. Stipa okmirii occurs exclusively in
Tajikistan (western Pamir; Dengubenko, 1980; Nobis,
2011, 2013; Gonzalo et al., 2012). Figure 4.
Notes. Stipa okmirii is likely a taxon of hybrid
origin. Because of its unigeniculate or indistinctly
bigeniculate awns, it could arise through hybridization
from S. lingua and either S. turkestanica or S. kirghi-
sorum (Nobis, 2013).
Stipa orientalis Trin. ex Ledeb., Fl. Altaic. [Ledebour]
1: 83. 1829. TYPE: [Kazakhstan.] In rupium fissuris
montis Arkaul, 17 May 1826, C. A. Meyer (lectotype,
designated by Tzvelev [1976: 583], LE!).
Within the species, two varieties are recognized.
1. Leaves of the vegetative shoots distinctly sca-
brous, ligules with hairs up to 1 mm
long...Stipa orientalis Trin. ex Ledeb. var. orientalis
19. Leaves of the vegetative shoots glabrous to slightly
scabrous, ligules with hairs 0.7–1.5(–2) mm
long.......................Stipa orientalis
var. azutavica (Kotukhov) M. Nobis & P. D. Gudkova
38 Annals of the
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Stipa orientalis var. orientalis.
Habitat. Stipa orientalis var. orientalis is found in
steppes, screes, rock crevices and ledges, and high
mountain deserts at 300–4500 m.
Distribution. The range of the taxon extends from
Iran (Alborz Mountains) up to southern Russia, Mongolia,
China, and northwest India (Tzvelev, 1968, 1976; Freitag,
1985; Wu & Phillips, 2006; Nobis et al., 2013). In Middle
Asia, it occurs in Afghanistan, China, Iran, Kazakhstan,
Kyrgyzstan, Pakistan, Tajikistan, and Uzbekistan.
Stipa orientalis var. azutavica (Kotukhov) M. Nobis &
P. D. Gudkova, Phytotaxa 245(1): 36. 2016. Basio-
nym: Stipa azutavica Kotukhov, Turczaninowia
1(2): 9–10. 1998. TYPE: Altaj australis, praemon-
tia australi-orientalia jugi Azutau, mons Bulgarta-
baty, desertum lapidosum, 22 May 1991, Ju.
Kotuchov s.n. (lectotype, designated by Nobis &
Gudkova [2016: 36], LE!; isolectotypes, KRA
436050!, KUZ!, LE!).
Habitat. Stipa orientalis var. azutavica is found in
rock crevices, ledges, and screes at 1000–1500 m.
Distribution. Stipa orientalis var. azutavica occurs
in the mountains of central Asia in Kazakhstan, Mon-
golia, Russia, and Pakistan (Kotukhov, 2002; Nobis &
Gudkova, 2016). In Middle Asia it is found in eastern
Kazakhstan (Sauro-Manrak Mountains) and Pakistan.
Notes. A third variety, Stipa orientalis var. lada-
khorum M. Nobis, is currently known only from north-
western India (Ladakh) but seems likely to extend into
Middle Asia. It differs from variety orientalis by its
vegetative shoots, which are densely pubescent versus
glabrous, respectively (Nobis & Nowak, 2016).
Stipa ovczinnikovii Roshev., Fl. URSS 2: 87. 1934.
TYPE: [Tajikistan.] Zeravshanskii khr, polynno-
kovyl’naya step po zap. sklonu gor, obrashchen-
nykh k doline Zeravshana, bliz kishlaka Syuzhena,
3 June 1932, P. Ovczinnikov 144 (holotype, LE!;
isotype, TAD 2342!).
Habitat. Stipa ovczinnikovii is found in steppe
grasslands at 1500–2200 m.
Distribution. Stipa ovczinnikovii is found in Tajiki-
stan (Hissar, Zeravshan, and Turkestan Mountains).
Notes. Stipa ovczinnikovii, with a Russian descrip-
tion, was published by Roshevitz (1934) with reference to
its Latin description in another publication. However, a
Latin description of the species was published only two
years later in 1936 [Trudy Bot. Inst. Acad. Nauk SSSR,
Fl. i Sist. Vyssh. Rast 1(2): 92. 1936].
Stipa pamirica Roshev., Bot. Mater. Gerb. Bot. Inst.
Komarova Akad. Nauk SSSR 11: 20. 1949. TYPE:
[USSR, Tajikistan.] Vakhan-Inkashimskii raion,
kovyl’no-solyankovo-polynnaya polupustynya po
vost. shchebnistym sklonam v raione kishlaka
Vrang, 3120 m, 9 Aug. 1935, P. Ovchinnikov &
K. Afanasev 1735 (lectotype, designated by Tzve-
lev [1976: 584], LE!). [Stipa badachschanica
Roshev. var. pamirica (Roshev.) M. Nobis.
Stipa parviflora Desf., Fl. Atlant. 1: 98–99, t. 29. 1798.
TYPE: In collibus aridis prope Mascar et in regno
Tunetano, Desfontaines (syntype, P not seen, after
Freitag, 1985). [Achnatherum parviflorum
(Desf.) M. Nobis, comb. nov. [[Stipellula parvi-
flora (Desf.) R¨oser & Hamasha, Schlechtendalia
24: 92. 2012].
Notes. Stipa parviflora is characterized by its
maize-like lemma epidermal pattern, which clearly
distinguishes it from other species of Stipa and
confirms that this species belongs to the achnathe-
roid group of grasses within the Stipeae. R¨oser
(2012) transferred this taxon to the new paraphyletic
genus Stipellula based on molecular analysis
(Hamasha et al., 2012). However, similar to Peter-
son et al. (2019), we recognize annual Stipellula
capensis as the only species in this genus originally
described by Tzvelev (1974, 2012), and prefer to
treat S. parviflora as a member of Achnatherum.How-
ever, it also has not been excluded that this annual
species mentioned above should be treated as a member
of Achnatherum (as A. capensis). Further studies using
modern molecular methods are needed to estabish a
phylogenetic relation within achnatheriod members of
the tribe Stipeae.
Stipa pavlovii Kotukhov, Turczaninowia 1(1): 7. 1998.
TYPE: Saur-Tarbagataj, jugi Saurici brachia
australi-occidentalia, 1800 m, in regione hiberna-
culi Kyzylkija, declivitas austro-orientalis saxosa,
4 July 1991, Ju. Kotukhov s.n. (holotype, LE not
found, Nobis & Gudkova, 2016). 5Stipa sczer-
bakovii Kotukhov.
Stipa pelliotii Danguy, Bull. Mus. Natl. Hist. Nat. 17:
451. 1911. TYPE: Asie Centrale, Zamutch-tagh,
1700 m, 28 July 1907, Pelliot & Vaillant 375
(holotype, P02241094!; isotype, LE!). [Achna-
therum pelliotii (Danguy) R¨oser & Hamasha, Pl.
Syst. Evol. 298: 365. 2012 [[Ptilagrostis pelliotii
(Danguy) Grubov, Consp. Fl. Outer Mongolia
(Vasc. Pl.) 62. 1955].
Stipa penicillata Hand.-Mazz., Oesterr. Bot. Z. 85:
226–227. 1936. TYPE: Kuku-nor, gegen Lombutong,
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Nobis et al. 39
Synopsis of Stipa (Poaceae) in Middle Asia

21 Sep. 1918, Licent 4869 (holotype, W1926-
0016157!; isotype, K00433026!).
Synonyms. 5Stipa laxiflora Keng.
Habitat. Stipa penicillata is found in steppes, high
mountain deserts, as well as rock crevices and ledges at
1500–4500 m.
Distribution. Stipa penicillata is found in China
(Tzvelev, 1968; Wu & Phillips, 2006). In Middle Asia,
it occurs in Tian-Shan, Kunlun, and Himalayas.
Notes. Leaf indumentum in this species is variable
(Wu & Phillips, 2006). Specimens with an abaxial surface
of scabrous vegetative leaves represent Stipa penicillata
var. penicillata, whereas specimens with vegetative leaves
and sheaths of culm leaves densely pubescent belong to S.
penicillata var. hirsuta P. C. Kuo & Y. H. Sun.
Stipa pennata L., Sp. Pl. 1: 78. 1753. TYPE: s. loc., A.
van Royen 900.320-437 (lectotype, designated by
Freitag [1985: 437], L!) (5Stipa joannis ˇ
Celak.).
Notes. The occurrence of the species in eastern
Kazakhstan reported by Kotukhov (2002) requires con-
firmation, whereas data given for Tajikistan from Gon-
zalo et al. (2013) refer, in fact, to Stipa kirghisorum not
S. pennata. Although we have seen no specimens of S.
pennata from Middle Asia, its presence in northern
Kazakhstan is probable.
Stipa pilgeriana K. S. Hao, Bot. Jahrb. Syst. 68(5): 583.
1938. TYPE: [China.] Kokonor, im Tsi-gi-gen-ben
Gebiete, 3900 m, 25 Aug. 1930, K. S. Hao 1009
(holotype, PEK not seen). 5Stipa purpurea
Griseb. (syn. after Freitag, 1985).
Stipa platypoda Bor, Biol. Skr. 14(4): 81. 1965. TYPE:
Afghanistan. Herat, profuse over the whole plain,
Aitchison 1137 (holotype, K!; isotype, BM!). 5
Stipa lingua A. Junge.
Stipa praecapillata Alechin, Krat. Predv. Otchet O
Rabotakh Nizhegorod Geobot. Eksped. 1925.
171. 1926. TYPE: Nizhegorodskaya gub., Lukya-
novskii u., bliz sel. Divii Usad, yuzhn. sklon
Kosovoi gory, 14 June 1925, V. Alekhin, K. Dobro-
khotova & I. Belov s.n. (lectotype, designated by
Tzvelev [1976: 581], LE!). 5Stipa sareptana
A. K. Becker.
Notes. Although we treat Stipa praecapillata as
conspecific with S. sareptana, it should not be excluded
that S. praecapillata (having a well-developed ring of
hairs at the lemma apex) is more closely related to S.
krylovii, from which it differs by having spinulose versus
glabrous vegetative leaves. Thus, taxonomic revision of
this group of taxa using modern molecular methods is
required.
Stipa 3pseudocapillata Roshev., Fl. Aziat. Ross. 12:
172, pl. 11, 6, 6a. 1916, pro sp. (S. sareptana A. K.
Becker 3S. lessingiana Trin. & Rupr.). TYPE: Stipa
consanguinea Trin.?, Songoria, Alex. Schrenk s.n.
(Herbarium Trautvetter) (holotype, LE!; isotype, LE!).
Synonyms. 5Stipa spiridonovii Roshev.
Habitat. Stipa 3pseudocapillata is found in sandy
and clayey semideserts and steppes at 300–700 m.
Distribution. Stipa 3pseudocapillata is found in
Middle Asia (Tzvelev, 1976), specifically in Kazakhstan
(Transcaspian and Balkhash regions).
Notes. Stipa 3pseudocapillata is regarded as a very
rare endemic species in the flora of Kazakhstan. In
2014 a new locality of this taxon was found in northern
Kazakhstan, Akmolinskii region, near Ereimentau
town, N part of Ereimentau Mountains, steppe, 27 June
2014, A. L. Ebel s.n. (det. M. Nobis, KRA). A specimen
of this taxon was collected from the locality where the
population of S. sareptana and S. lessingiana grow
together. Tzvelev (1976) suggested that S. pseudocapil-
lata is a species of hybrid origin and indicated S.
capillata and S. lessingiana as the parental species.
In our opinion, hybridization between S. sareptana and
S. lessingiana is more probable than between S. capil-
lata and S. lessingiana because these two first species
flower in the same period (usually in May), while S.
capillata flowers much later (usually June and July).
Stipa 3pseudocapillata was described by Roshevitz
(1916) based on one gathering and two herbarium sheets
preserved at LE. Tzvelev (1976) stated that the holotype
of this species is preserved at LE, but at the same time
he did not mention the second sheet of this species that
was also preserved at LE. Because only one of them has
the stamp “Herbarium Trautvetter,”as was mentioned
by Roshevitz (1916) in the protologue of the taxon, it
should be treated as the holotype. Taking into account
that the herbarium sheet with the holotype of
S. 3pseudocapillata has no number, for identification
purposes the following information is useful: on the
sheet, the two additional illegible labels written in
German were attached to Roshevitz’s determination
label: “Stipa pseudocapillata Roshev., I 1916, determ.
R. Roshevitz”and two labels (white and blue) with the
word Typus.
Stipa 3pseudomacroglossa M. Nobis, nothosp. nov.
(S. lipskyi Roshev. 3S. macroglossa P. A. Smirn.
subsp. macroglossa). TYPE: Dolina rzeki Iskan-
derdarya (Iskanderdarya River valley) Rejon Zer-
awsza´nski B (Zeravshan Region B), Zachodni
40 Annals of the
Missouri Botanical Garden

Tad˙zykistan (West Tajikistan) Pamiro-Alai; Mur-
awy stepowe w´sr´od głaz ´ow, na stokach g ´or, na
prawym zboczu doliny rzeki Iskanderdarya, ok.
1.5 km na E od Jeziora Iskanderkul (steppe grass-
lands, among stones, on mtn. slopes, on the right
slope of Iskanderdarya River valley, ca. 1.5 km of
Iskanderkul Lake), 39°049N / 68°239E, 2450 m,
10 June 2012, M. Nobis s.n. (holotype, KRA
476856!; isotypes, KRA 479081!, KRA 479082!).
Figure 10.
Plants perennial, densely tufted, with a few culms
and numerous vegetative shoots; culms 35–45 cm tall,
3-noded, glabrous at nodes and shortly pubescent below
them. Leaves of vegetative shoots: sheaths glabrous,
with white edge at margins ciliate; ligules rounded or
acute, (0.2–)0.3–1.4(–2) mm and ciliate at margins with
cilae (0.4–)0.8–1.4 mm; blades of vegetative shoots
convolute, 11–14 cm, 0.4–0.6 mm in diam., upper
surface densely short-pilose with hairs 0.2–0.3 mm,
glabrous and smooth beneath, sporadically somewhat
scabrous due to short, single prickles, young leaves with
tuft of hairs at apex. Cauline leaves: sheaths glabrous,
with white and glabrous margins, shorter than inter-
nodes, upper sheath up to 15 cm, glabrous, in upper part
Figure 10. Stipa 3pseudomacroglossa M. Nobis. —A. Holotype. —B. Top of lemma. —C. Lemma micromorphological
pattern. —D. Callus. —E. Adaxial surface of the vegetative leaf. —F. Abaxial surface of the vegetative leaf. Scale bars: A 55 cm,
B, E, F 50.3 mm, C 530 mm, D 51 mm.
Volume 105, Number 1
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Nobis et al. 41
Synopsis of Stipa (Poaceae) in Middle Asia

scabrous, slightly inflated; ligules up to 2.3 mm, trun-
cate or acute, ciliate at apex; blades glabrous to slightly
scabrous, 4–7 cm. Panicle 14–20 cm contracted, at base
enclosed by sheath of uppermost leaf, branches erect,
setulose, single or paired. Glumes subequal, 44–52 mm,
narrowly lanceolate, tapering into long hyaline apex,
midvein sometimes setulose with cilia up to 1 mm.
Anthecium 13–14.5 30.8–1 mm. Callus 1.9–2.4 mm,
densely long-pilose, base of callus not enlarged, periph-
eral ring ca. 0.3 mm in diam., acute, cuneate, scar broadly
elliptic. Lemma pale green, on dorsal surface with abun-
dant hooks and 7 lines of ascending hairs, up to 0.7 mm,
ventral lines terminating at 1–1.5 mm below top of lemma,
dorsal line terminating at 2.5–3.5 mm below top of lemma;
top of lemma glabrous (without a ring of hairs) and
scabrous due to short prickles below apex. Palea equal
to lemma in length. Awn 158–220 mm, unigeniculate or
indistinctly bigeniculate; column 38–45 mm, twisted,
0.4–0.5 mm wide at base, glabrous and smooth; seta
slightly flexuous 120–176 mm and 2.5–3.5 times longer
than column, hairs in lower part of seta 5.2–6.2 mm,
gradually decreasing in length toward apex. Anthers yel-
low, 8–9 mm, glabrous.
Phenology. Stipa 3pseudomacroglossa flowers from
May to June.
Habitat. Stipa 3pseudomacroglossa is found in
mountain steppes at 2350–2500 m.
Distribution. Stipa 3pseudomacroglossa occurs in
Tajikistan (Zeravshan Mountains). Figure 9.
Stipa pulcherrima K. Koch, Linnaea 21: 440. 1848.
TYPE: Central Poland, Nida Basin, western part of
gypsum hills, located betw. Bronina & Owczary
villages near Busko-Zdr´oj, 50°279N / 20°449E, 23
May 2010, A. Nobis & M. Nobis s.n. (holotype,
KRA 465416!; isotypes, BM!, E!, GOET!, KRA
465417!, PR!, WA!), typ. cons. prop. by Nobis
et al. (2017b: 518).
Stipa pulcherrima subsp. pulcherrima.
Synonyms. [Stipa pennata L. subsp. pulcherrima
(K. Koch) ´
A. L¨ove & D. L¨ove, Folia Geobot. Phytotax.
10(3): 273. 1975; [S. pennata subsp. pulcherrima (K.
Koch) Freitag, Notes Roy. Bot. Gard. Edinburgh 42(3):
440. 1985; [S. pennata var. pulcherrima (K. Koch)
Hal´acsy, Consp. Fl. Graec. 3: 352. 1904; [S. pennata
var. pulcherrima (K. Koch) Beck, Wiss. Mitt. Bosnien &
Herzegovina 9: 426. 1904; [S. pennata fo. pulcherrima
(K. Koch) Brand, Syn. Deut. Schweiz. Fl. 3: 2718.
1907; 5S. grafiana Steven; [S. pulcherrima K. Koch
subsp. grafiana (Steven) Pacz., Khersonsk. Fl. 1: 115.
1914; [S. pennata L. var. grafiana (Steven) Lindem.,
Fl. Chers. 2: 283. 1882.
Habitat. Stipa pulcherrima subsp. pulcherrima is
found in mountain steppes, clay slopes, and grasslands
at ca. 400–2400 m.
Distribution. Stipa pulcherrima subsp. pulcherrima
is widely distributed; its range extends from central
Europe and the Mediterranean area, through the Cau-
casus, Black Sea region, southern Ural to Siberia (Altai
Mountains), and central Tian-Shan Mountains in Mid-
dle Asia (Tzvelev, 1976, 2006; Martinovsk´y, 1980;
Freitag, 1985; Durka et al., 2013; Gudkova et al.,
2014b). In Middle Asia, it occurs in Kazakhstan and
Kyrgyzstan (central Tian-Shan Mountains).
Notes. This species was confirmed in the studied
area in Kyrgyzstan (central Tian-Shan, ca. 36.5 km NNE
of Naryn, ca. 63 km NE of At-Bashy, steppe on slope/
shrubs, 41°299410N / 76°259330E, elev. 2283 m, exp.
N, incl. 25°, 6 July 2018, M. Nobis, E. Klichowska, A.
Wr´obel & A. Nowak s.n. [KRA 0502537, KRA 0502538,
KRA 0502540, KRA 0502541, KRA 0502542, KRA
0502544]) and Kazakhstan (63 km NW of Usharal, E of
Balkhash Lake, steppe, 46°319380N / 80°179230E,
elev. 404 m, 23 May 2019, M. Nobis s.n. [KRA]); up
to now, its occurrence in Middle Asia was questionable
(Pazij, 1968; Tzvelev, 1976).
Because the lectotype of Stipa pulcherrima desig-
nated by Freitag (1985: 440) (Tatarisch Grusien, 1844,
K. Koch s.n. [GOET No. 013762]) creates serious no-
menclatural conflict (Nobis et al., 2017b), as it can be
referable to S. endotricha Martinovsk´y, S. majalis Klo-
kov, or S. pulcherrima var. alagezica Tzvelev, Nobis
et al. (2017b) proposed a gathering from central Europe
as the conserved type. This gathering demonstrates
characters that are fully within the range of variability
of S. pulcherrima s. str. as well as occurring in spec-
imens from Georgia or Armenia (Transcaucasia, Tiflis,
in fruticetis, [19]20–V–20, A. Grossheim s.n. [LE]), from
where the species was described.
Stipa pulcherrima subsp. crassiculmis (P. A. Smirn.)
Tzvelev, Novosti Sist. Vyssh. Rast. 11: 18. 1974.
Basionym: Stipa crassiculmis P. A. Smirn., Repert.
Spec. Nov. Regni Veg. Beih. 22: 375. 1926.
Habitat. Stipa pulcherrima subsp. crassiculmis is
found in steppe grasslands at 800–1500 m.
Distribution. Stipa pulcherrima subsp. crassiculmis
is found in eastern Europe and southwestern Asia
(Tzvelev, 1976; Martinovsk´y, 1980; Nikitin & Geldi-
khanov, 1988). In Middle Asia, it occurs in northern
Iran and Turkmenistan (Kopet-Dagh Mountains).
Stipa purpurascens Hitchc., Proc. Biol. Soc. Washington
43: 95. 1930. TYPE: [China.] Gansu, S of Sining,
in the La Che Tze Mtns., 3350–3900 m, Aug.
42 Annals of the
Missouri Botanical Garden

1923, R. C. Ching 686 (holotype, US-1245701 not
seen). [Stipa regeliana Hack.
Stipa purpurea Griseb., Nachr. K¨onigl. Ges. Wiss.
Georg-Augusts-Univ. 3: 82. 1868. TYPE: [China.]
Tibet, Gnari (Nari) Khorsum, 5000 m, 5–15 Sep.
1855, Schlagnitweit 7116 (holotype, GOET!; iso-
types, BM!, LE!).
Synonyms. [Ptilagrostis purpurea (Griseb.) Roshev.,
Fl. URSS 2: 76. 1934; 5Lasiagrostis tremula Rupr.,
Ost.-Sack. & Rupr., M´em. Acad. Imp., St. P ´etersb., S ´er.
7, 14(4): 35. 1869. TYPE: Fl. Sarymeki, sudlicher
Abhang des Tian-Schan, 28 July 1867, F. Osten-
Sacken s.n. (holotype, LE!); 5Ptilagrostis semenovi
Krasn., Spisok rastenii sobrannykh v vostochnom
Tyan-Shane, letom 1886 goda, 125, 1887. TYPE: Ad
fl. Sary-Jassy, 1 Aug. 1886, Krassnov s.n. (holotype, LE);
[Stipa semenovi Krasn.
Habitat. Stipa purpurea is found in alpine meadows
at 2500–4700 m.
Distribution. Stipa purpurea is found in the moun-
tains of central Asia (from Tian-Shan up to the Hima-
layas; Tzvelev, 1968, 1976; Freitag, 1985; Wu &
Phillips, 2006). In Middle Asia, it occurs in eastern
Afghanistan, western and southwestern China, eastern
Kyrgyzstan, eastern Tajikistan, and eastern Pakistan.
Notes. This taxon is highly variable in morphology,
and its taxonomic revision is required.
Stipa regeliana Hack., Sitzungsber. Kaiserl. Akad.
Wiss., Math.-Naturwiss. Cl., Abt. 1, 89: 130. 1884.
TYPE: [Kyrgyzstan.] Issyk-Kul, Musart, 7–50009,
Aug. 1877, A. Regel s.n. (holotype, W 1916-
0026455!).
Synonyms. [Achnatherum regelianum (Hack.)
Tzvelev, Novosti Sist. Vyssh. Rast. 43: 22. 2012; 5
Stipa purpurascens Hitchc.
Habitat. Stipa regeliana is found in alpine meadows
at 2200–4500 m.
Distribution. Stipa regeliana is a widely distributed
central Asian species (Pazij, 1968; Tzvelev, 1968,
1976; Freitag, 1985; Wu & Phillips, 2006; Nobis
et al., 2015b). In Middle Asia, it occurs in eastern
Afghanistan, western and southwestern China, eastern
Kazakhstan, eastern Kyrgyzstan, eastern Tajikistan,
and northeastern Pakistan.
Stipa retorta Cav.,Observ.Hist.Nat.1:119.1795.TYPE:
MA not seen. 5Stipellula capensis (Thunb.) R¨oser
& Hamasha, Schlechtendalia 24: 92. 2012. [[Stipa
capensis Thunb., syn. after Freitag, 1985].
Stipa richteriana Kar. & Kir., Bull. Soc. Imp. Natural-
istes Moscou 14(4): 862. 1841. TYPE: [Eastern
Kazakhstan.] In lapidosis mont. Arganty, 1840,
Karelin 907 (lectotype, designated here, LE!; iso-
lectotype, LE!).
Notes. There are six sheets in LE with plants rep-
resenting syntypes of Stipa richteriana. One of these
sheets, labeled “Stipa richteriana Kar. et Kir. enum.
1840, no. 907, in lepidosis mont. Arganatty, 1842 Jul,
Kraelin,”has an additional label attached by N. Tzve-
lev: “Stipa richteriana Kar. et Kir., Lectotype, 1972.”
However, in his work Tzvelev (1976) indicated it as a
type (holotype) and stated that there are three additional
isotypes preserved at LE. In accordance with the Inter-
national Code of Nomenclature, Tzvelev’s typification
does not meet the requirements of effective lectotypi-
fication (McNeill et al., 2012; Turland et al., 2018).
Since the collection of typical specimens of S. richteri-
ana represents a collection of syntypes, the lectotype
should be designated. Of the two sheets with specimens
of S. richteriana and the original label, dated 1840 and
signed/collected by Karelin, we designate the one,
previously selected by Tzvelev in 1972, as the lectotype
and the second as isolectotype. The other four speci-
mens are syntypes.
Stipa richteriana subsp. richteriana.
Within the taxon two varieties are recognized.
1. Leaves of vegetative shoots glabrous to slightly
scabrous . . . Stipa richteriana Kar. & Kir. var. richteriana
19. Leaves of vegetative shoots densely pubescent . . .
... Stipa richteriana var. dasyphylla (Roshev.) Tzvelev
Stipa richteriana var. richteriana.
Synonyms. 5Stipa kuhitangi Drobow; 5S. vor-
onini Krasn.; [S. woroninii Krasn.
Habitat. Stipa richteriana var. richteriana is found
in steppes, stony and clay slopes, and sandy grasslands
at 200–1200 m.
Distribution. Stipa richteriana var. richteriana is
found in MiddleAsia and southwestern Siberia (Lavrenko
& Nikolskaya, 1965; Tzvelev, 1976; Nobis et al., 2016b).
In Middle Asia it occurs in western China, Kazakhstan,
and Uzbekistan.
Stipa richteriana var. dasyphylla (Roshev.) Tzvelev,
Zlaki SSSR 578. 1976. Basionym: Stipa richteri-
ana f. dasyphylla Roshev., Fl. Aziat. Ross. 12:
135. 1916. TYPE: Fl. Iliensis, prope Andrakai,
May 1886, Krassnov s.n. (lectotype, designated by
Tzvelev [1976: 578], LE!).
Volume 105, Number 1
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Nobis et al. 43
Synopsis of Stipa (Poaceae) in Middle Asia

Habitat. Stipa richteriana var. dasyphylla is found
in steppes, stony and clay slopes, and sandy grasslands
at 200–1200 m.
Distribution. In Middle Asia, Stipa richteriana var.
dasyphylla is found in central Kazakhstan, within the
range of the nominal variety (Tzvelev, 1976).
Stipa richteriana subsp. jagnobica (Ovcz. & Czu-
kav.) Tzvelev, Novosti Sist. Vyssh. Rast. 11: 14.
1974. Basionym: Stipa jagnobica Ovcz. & Czukav.,
Izvest. Otdel. Estestven. Nauk Akad. Nauk Ta-
dzhik SSR 17: 51. 1957.
Within the taxon two varieties are recognized.
1. Leaves of vegetative shoots glabrous . . .........
.... Stipa richteriana Kar. & Kir. var. jagnobica
(Ovcz. & Czukav.) M. Nobis & A. Nowak
19. Leaves of vegetative shoots densely pubescent . . .
Stipa richteriana var. hirtifolia M. Nobis & A. Nowak
Stipa richteriana var. hirtifolia M. Nobis & A.
Nowak, var. nov. TYPE: Tajikistan. Dolina rzeki
Archamaidon (Archamaydon River valley) Rejon
Zerawsza´nski B (Zeravshan Region B), Zachodni
Tad˙zykistan (western Tajikistan) Pamir Alai Mtns.;
wysokog´orska mrawa stepowa (step bylicowo-
ostnicowy), na zachodnich stokach g´or, na prawym
zboczu doliny rzeki Archamaidon, na S od wsi
Gazza koło Zimtut [high mtn. steppe grassland, on
western mtn. slopes, on the right slope of the
Archamaydon River valley, S of Gazza settlement
near Zimtut], 39°149440N / 68°019550E, 1740 m,
exp. W, slope 45°, pH 58.2, 16 June 2010, M.
Nobis s.n. (holotype, KRA 468742!; isotypes, KRA
468760!, KRA 468750!, KRA 465273!).
Diagnosis. Stipa richteriana Kar. & Kir. var. hirtifolia M.
Nobis & A. Nowak differs from S. richteriana var. jagnobica (Ovcz.
& Czukav.) M. Nobis & A. Nowak in having the abaxial surface of
the vegetative leaves densely pubescent versus the abaxial surface
of the vegetative leaves glabrous or slightly scabrous.
Habitat. Stipa richteriana var. hirtifolia is found
in high mountain steppes at 1700–2400 m.
Distribution. In Middle Asia, Stipa richteriana var.
hirtifolia is found in Tajikistan, within the range of the
nominal variety (Zeravshan Mountains).
Paratypes. TAJIKISTAN. Fan River valley, Zeravshan
Region C, western Tajikistan, Pamir Alai Mtns.; high mtn.
steppe grassland, on western mtn. slopes, Gushty Mtns., E of
Zeravshan I settlement S of Aini, 39°139N / 68°330E, 2400 m,
exp. W, slope 35°, 19 June 2010, M. Nobis s.n. (KRA 468744,
KRA 468743).
Stipa richteriana var. jagnobica (Ovcz. & Czukav.)
M. Nobis & A. Nowak, comb. & stat. nov. Basio-
nym: Stipa jagnobica Ovcz. & Czukav., Izvest.
Otdel. Estestven. Nauk Akad. Nauk Tadzhik.
SSR 17: 51. 1957.
Habitat. Stipa richteriana var. jagnobica is found in
steppes, stony and clay slopes, and rocky grasslands at
1700–3000 m.
Distribution. Stipa richteriana var. jagnobica is
found in Tajikistan (Hissar and Zeravshan Mountains;
Ovchinnikov & Chukavina, 1957) and in central-
eastern Afghanistan (Freitag, 1985). Figure 4.
Stipa roborowskyi Roshev., Bot. Mater. Gerb. Glavn.
Bot. Sada R.S.F.S.R. 1(6): 1. 1920. TYPE: Kuen-
Lun, severn. sklon khrebta Russkogo, 10–12.5 tys.
fut. abs. vysoty, ur. Kara-sai, Lyundzhilin-
Khakum, verkhovya reki Aksu, po lessovym sklo-
nam gor, luga, 3 July 1890, W. J. Roborowski s.n.
(holotype, LE!; isotype, W 1916-0034934!).
Habitat. Stipa roborowskyi is found in high moun-
tain deserts, alpine meadows, and stony slopes at
3000–4500 m.
Distribution. Stipa roborowskyi is found in the
mountains of central Asia: Pamir, Karakorum, Kunlun,
and Himalayas (Wu & Wang, 1999; Nobis et al.,
2014b). In Middle Asia it occurs in western China
and northwestern Pakistan.
Notes. Stipa roborowskyi is morphologically similar
to S. klimesii M. Nobis [syn. [Stipa basiplumosa Munro
ex Hook. f. var. longearistata Munro ex Hook. f.], which
occurs in China (Tibet), Bhutan, Nepal, India (Ladakh),
and Pakistan; however, S. roborowskyi differs from S.
klimesii by having shorter ligules of the vegetative shoots
(0.5–1.5[–2] vs. [2–]3.5–7.5[–9] mm), shorter anthecium
([6–]6.5–7.5[–7.7] vs. [7–]8.3–9.5[–10] mm), and shorter
hairs on the seta ([0.3–]0.5–1.1[–1.4] vs. [1–]1.3–2
[–2.3] mm).
New record of Stipa roborowskyi to the flora of Pakistan.
Karakorum, Upper Braldu tributary, above Chikor, 36°22–249N
/ 75°22–249E, alpine steppe, alt. 4100–4200 m, 17 Aug. 1991,
G. & S. Miehe 6221 (MSB).
Stipa roerichii Keng, J. Wash. Acad. Sci. 28: 307. 1938.
TYPE: Mongolia. In crevice of exposed rocks,
Temur Khada, Peiling Miao, Suiyan Prov.,
1500 m, 26 July 1935, Y. L. Keng 3181 (Roerich
Exp. no. 518) (holotype, Herb. of the Nat. Research
Inst. of Biol., Acad. Sinica, Nanking, China not
seen). 5Achnatherum nakai (Honda) Tateoka
ex Imzab, Fl. Intramongol 7: 196. 1983 [[Stipa
nakaii Honda] (syn. after Wu & Phillips, 2006).
Stipa rubens P. A. Smirn., Repert. Spec. Nov. Regni
Veg. 21: 231. 1925. TYPE: Akmolinskaya obl. i u.,
na stepennoi polyane v berezovom lesu v 11
44 Annals of the
Missouri Botanical Garden

verstakh k vost. ot Akmolinska, 27 May 1914, S.
Ganeshin 351 (holotype, LE!; isotypes, LE!,
MW!). 5Stipa zalesskii Wilensky var. zalesskii.
Stipa rubentiformis P. A. Smirn., Trudy Imp. S.-
Peterburgsk. Bot. Sada 40: 115. 1928. TYPE:
Saratov, okr. Polivanovka, stepnoi uchastok, 1 June
1927, B. A. Fedchenko & E. G. Bobrov 30 (lectotype,
selected by Tzvelev in 1970 and designated here,
LE!). 5Stipa zalesskii Wilensky var. zalesskii.
Stipa sabulosa (Pacz.) Sljuss., Trudy Nauchno-Issl. Inst.
Bot. 37: 26. 1963. Basionym: Stipa pennata L.
subsp. joannis ˇ
Celak. f. sabulosa Pacz., Khersonsk.
Fl. 1: 112. 1914. TYPE: Ukraine. Lugov forest
country house, Tyasmin, Aleks, u., 18 May 1911, I.
Paczoski (lectotype, designated by Tzvelev [1976:
591], LE!). [Stipa borysthenica Klokov ex
Prokudin var. borysthenica.
Stipa saikanica Kotukhov, Turczaninowia 1(2): 10.
1998. TYPE: Saur-Tarbagataj, praemontia boreali-
occidentalia jugi Saikan, locus Akseir, denudationes
argillarum tertiariarum (in gypsaceis), partitiones
glareoso-argillosae, 9 June 1992, Ju. Kotuchov
s.n. (lectotype, designated by Nobis & Gudkova
[2016: 37], LE!; isolectotypes, KRA 436036!,
KUZ! [2 sheets], LE! [2 sheets]). 5Stipa les-
singiana Trin. & Rupr.
Stipa saposhnikowii (Roshev.) Kitag., Rep. Inst. Sci.Res.
Manchoukuo 6(4): 118. 1942. [Achnatherum
saposhnikowii (Roshev.) Nevski, Trudy Bot. Inst.
Akad. Nauk S.S.S.R., Ser. 1, Fl. Sist. Vyssh. Rast. 4:
224. 1937. Basionym: Timouria saposhnikowii
Roshev., Fl. Aziat. Ross. 12: 174. 1916. TYPE:
Semirech. obl., Przhev. u., u. R. Sary-Dzhas pri uste
Kaindy, kamenistye sklony k reke i priberezhnaya
galka, 8 Aug. 1912, V. Sapozhnikov & B. Shishkin
s.n. (lectotype, designated here, LE! [sheet with a
single specimen]; isolectotype, LE!).
Stipa sareptana A. K. Becker, Bull. Soc. Imp. Natural-
istes Moscou 57(1): 52. 1882. TYPE: Sarepta,
1881, A. Becker 3887 (lectotype, designated here,
LE!).
Synonyms. [Stipa capillata L. var. sareptana (A.
K. Becker) Schmalh., Fl. Sr. i Yuzh. Ross. 2: 595. 1897;
5S. akseirica Kotukhov; 5S. sareptana A. K. Becker
var. kasakorum Roshev., Fl. URSS 2: 111. 1934. TYPE:
LE not seen; (?) 5S. praecapillata Alechin; [S.
sareptana fo. praecapillata (Alechin) Krylov ex Roshev.,
Fl. URSS 2: 111. 1934.
Habitat. Stipa sareptana is found in steppes, stony
slopes, roadsides, escarpments, and fallows at 300–2500 m.
Distribution. Stipa sareptana is a widely distributed,
eastern European–central Asian species, its range
extending from southwestern Russia and Middle
Asia, up to Mongolia and western China (Tzvelev,
1976; Wu & Phillips, 2006; Gudkova et al., 2017a).
In Middle Asia, it occurs in western China, Kazakhstan,
Kyrgyzstan, Tajikistan, Turkmenistan, and Uzbekistan.
Figure 8.
Notes. Stipa sareptana was described by Becker
(1882) based on plant material collected from Sarepta
(currently part of Volgograd [southwestern Russia])
with a short diagnosis in German and without a date
of collection, collection number, or name of the herbar-
ium where the type was deposited. During revision of the
material representing S. sareptana that was preserved at
LE, W, and WU, we found 12 potential syntypes. All
had been collected by A. Becker from Sarepta.
However, a part of them have either no collection date
or different collection numbers and descriptions on the
labels. Tzvelev (1976) mentioned that the type and four
isotypes of S. sareptana are preserved at LE. However,
we found another five specimens (without year of col-
lection) labeled by L. Pignotti in 2011 as a type of the
species (S ¨ud Russland, Sarepta, A. Becker [W-Hackel
1916-0022319]) in the herbarium of W as well as one
specimen labeled by M. Sonnleitner in 2013 as a type of
the species (Sarepta, 1881, A. Becker [WU 0072643!])
in WU. In accordance with the International Code of
Nomenclature (McNeill et al., 2012; Turland et al.,
2018), all of the specimens from Sarepta collected by
Becker and deposited at LE, W, and WU should be
treated as syntypes, and a lectotype needs to be des-
ignated. As a lectotype we designated the specimen
preserved at LE with the labels: “Stipa capillata L.,
Stipa sareptana Becker, Sarepta, 1881, A. Becker and
collection no. 3887,”with the additional label below
having a pencil-written description of morphological
characters as well as Smirnov’s revision label “Stipa
sareptana Becker, 1928.I, Det. P.A. Smirnow.”The other
specimens, attached to the five sheets preserved at LE,
five preserved at W, and one at WU, are the syntypes.
Stipa saurica Kotukhov, Bot. Zhurn. (Moscow & Lenin-
grad) 79: 103. 1994. TYPE: Saur-Tarbagatai, bra-
chia boreali-occidentalia jugi Saur, in viciniis
hibernaculi Kesek, clivulus saxosus australi-
occidentalis, in schistosis, 14 July 1992, Ju.
Kotuchov s.n. (lectotype, designated by Gonzalo
et al. [2011: 433], LE!; isolectotype, KUZ!). 5
Stipa 3manrakica Kotukhov.
Stipa sczerbakovii Kotukhov, Bot. Zhurn. (Moscow &
Leningrad) 76(6): 872. 1991. TYPE: Altai Aus-
tralis, brachia austro-orientalia jugi Azutau,
montes Bulgartabaty, elev. 600 m, locus Sargalym,
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Nobis et al. 45
Synopsis of Stipa (Poaceae) in Middle Asia

steppa fruticeto-graminosa lapidosa, 16 June
1988, J. Kotuchov s.n. (holotype, LE!; isotypes,
KRA 136031!, KUZ! [13 sheets]).
Synonyms. 5Stipa kyzylkiensis Kotukhov; 5S.
pavlovii Kotukhov.
Habitat. Stipa sczerbakovii is found in steppe grass-
lands at 600–1700 m.
Distribution. Stipa sczerbakovii is found in Kazakh-
stan, western China, Mongolia, and Russia (Kotukhov,
2002; Nobis & Gudkova, 2016; Zhao et al., 2019). In
Middle Asia, it occurs in eastern Kazakhstan (Tarba-
gatai Mountains).
Notes. Stipa sczerbakovii likely has a hybrid origin
(S. krylovii 3S. orientalis). The variability of morpho-
logical characters of this taxon is greater than that
presented by Kotukhov (1991) in the protologue: e.g.,
ligules of vegetative shoots (0.1–)0.3–2.7 mm (not 0.3–2mm
as in the protologue), anthecium 8–10.5(–11) mm (not
8.5–9mm),awns8.5–11.7 cm (not 9–10 cm), hairs on
seta 0.4–1.3 mm (not 0.5–0.8 mm), anthers 3.2–4.3 mm (not
3.5–4 mm). Taking into account that Kotukhov described S.
monticola (nom. nov. S. kotuchovii) as very similar to S.
sczerbakovii, taxonomic revision of the species using modern
molecular methods is required.
Stipa semenowi Krasn., Scripta Bot. Horti Univ. Imper.
Petropolitanae, Botanicheskiia Zapiski 2(1): 22.
1887. TYPE: Ptilagrostis semenovii, Ad fl. Sary-
Jassy, 1 Aug. 1886, Krassnov s.n. (holotype, LE). 5
Stipa purpurea Griseb.
Notes. This taxon was also described by Krassnov
as Stipa semenovi Krasn. (Zap. Imperat. Russ. Geograf.
Obsch., Opyt istor. rasv. fl. yuzhn. ch. vost. Tyan-
Shanya 19: 341–342. 1888).
Stipa sibirica (L.) Lam., Tabl. Encycl. 1: 158. 1791.
Basionym: Avena sibirica L., Sp. Pl. 1: 79. 1753.
TYPE: Siberia, Amman 27 (lectotype, designated
by Scholz in Cafferty et al. [2000: 248], LINN-
95.1). 5Achnatherum sibiricum (L.) Keng ex
Tzvelev, Probl. Ekol. Geobot. Bot. Geogr. Florist.
140. 1977.
Stipa smirnovii Martinovsk´y, Preslia 47(3): 260. 1975.
TYPE: Bohemia, bor.–occ. Montges ˇ
Cesk´e
stˇredohoˇr´ı: in stepposis ad declivem austro occ.
Collies Ran´a, June 1974, Martinovsk´ys.n.(holotype,
PRC!). 5Stipa zalesskii Wilensky var. zalesskii.
Stipa spiridonovii Roshev., Bull. Jard. Bot. Acad. Sci.
URSS 1931, 30: 302. 1932. TYPE: Mangyshlak, v
1 km ot kotlov. Khangababa po doroge v fort Alek-
sandrovsk, pologii yuzhn. sklon ravniny sarmata,
suglinok, 8 June 1926, M. Spiridonov 755/9 (holo-
type, LE!). 5Stipa 3pseudocapillata Roshev.
Stipa splendens Trin., Neue Entdeck. Pflanzenk. 2: 54.
1821. TYPE: [USSR, Transbaikalia] Agrostis long-
iaristata, herb. Fischer (holotype, LE TRIN!). [
Neotrinia splendens (Trin.) M. Nobis, P. D.
Gudkova & A. Nowak, Turczaninowia 22(1): 40.
2019 ([Achnatherum splendens (Trin.) Nevski,
Trudy Bot. Inst. Akad. Nauk S.S.S.R., Ser. 1, Fl.
Sist. Vyssh. Rast. 4: 224. 1937).
Stipa staintonii Bor, Bull. Bot. Surv. India 7: 133. 1965.
TYPE: Nepal. Near Seng Khola, 12,500 ft. [3810
m], exposed cliffs, 4 Oct. 1954, Stainton, Sykes &
Williams 4677 (holotype, K!; isotype, BM!). [
Achnatherum staintonii (Bor) M. Nobis & P.
D. Gudkova, PhytoKeys 128: 112. 2019 ([Stip-
ella staintonii (Bor) R¨oser & Hamasha, Pl. Syst.
Evol. 298: 365. 2012, nom. illeg. [Stipellula
staintonii (Bor) R¨oser & Hamasha, Schlechtenda-
lia 24: 92. 2012).
Stipa stapfii Roshev., Bot. Mater. Gerb. Glavn. Bot. Sada
R.S.F.S.R. 5: 11. 1924. TYPE: Persia. [Fars Prov.,
near Shiraz], 1885, Stapf [1530] (holotype, LE!;
isotypes, WU072646!, WU072647!). 5Stipa
3assyriaca Hand.-Mazz.
Stipa subbarbata B. Keller, Bot.-Geogr. Issledov. Zai-
sansk. uj. Semipalatinsk. Obl. 2: 53. 1912. TYPE:
Kaldzhirskaya dolina, po levuyu stronu reki Kaldz-
hira, Chiganchii protoki v podgornom stepii, 4 July
1908, B. A. Keller s.n. (holotype, LE!; isotype,
LE!). 5Stipa hohenackeriana Trin. & Rupr.
Stipa 3subdrobovii M. Nobis & A. Nowak, nothosp.
nov. (S. drobovii (Tzvelev) Czerep.3S. caucasica
Schmalh.). TYPE: Dolina rzeki Iskanderdarya
(Iskanderdarya River valley) Rejon Zerawsza ´nski
B (Zeravshan Region B), Zachodni Tad˙zykistan
(western Tajikistan) Pamir Alai Mtns.; Zeravshan
Mtns., high mtn. steppe, betw. stones, on the left
slope of the Iskanderdarya River valley, ca. 0.5 km
E of Serimadarun Lake (near Iskanderkul Lake),
39°059N / 68°239E, 2340 m, inclination W, slope
5°, 15 June 2012, no. 3, M. Nobis & A. Nowak s.n.
(holotype, KRA 456386!). Figure 11.
Diagnosis. Stipa 3subdrobovii M. Nobis & A. Nowak is
most similar to S. drobovii (Tzvelev) Czerep., but differs in its
longer callus, (1.5–)1.6–2.1(–2.3) mm versus 0.8–1.3(–1.4)
mm, flexuous to straight versus strongly falcate hairs at the
dorsal part of the callus as well as shorter hairs at the adaxial
surface of leaves (0.15–0.25 vs. 0.35–0.6 mm). Stipa 3sub-
drobovii is also somewhat similar to S. caucasica var. fanica, but
differs in its somewhat shorter anthecium with lines of hairs on
the lemma reaching the top of the lemma or terminating
46 Annals of the
Missouri Botanical Garden

0.5–1 mm below the top of the lemma versus lines of hairs on
the lemma terminating 2–5 mm below the top of lemma, hairs in
the dorsal part of the callus slightly flexuous and as long as
those in the ventral part versus hairs in the dorsal part of the
callus straight and 3 to 4 times shorter than those in the ventral
part.
Plants perennial, densely tufted, with a few culms
and numerous vegetative shoots; culms (25–)30–60 cm
tall, 3-noded, pubescent or glabrous at nodes and
densely pubescent below them. Culm leaves: sheaths
glabrous or the lower shortly pubescent, blades glabrous
at abaxial surface and pubescent at adaxial surface.
Leaves of vegetative shoots: ligules up to 0.2 mm long,
densely up to 1.5 mm long, ciliate; abaxial surface of
blades glabrous and smooth or pubescent, adaxial sur-
face densely covered with hairs 0.15–0.25(–0.4) mm.
Glumes 35–46 mm. Anthecium 9–11 30.8–1.1 mm.
Callus acute, (1.5–)1.6–2.1(–2.3) mm, its base 0.7–0.9 3
0.25–0.3 mm, densely bearded, dorsal hairs flexuous or
straight. Lemma with 7 lines of hairs, marginal and dorsal
lines reaching base of awn or terminating at 0.2–1mm
below top; lemma apex with ring of hairs. Awn unigeni-
culate, column 20–35 mm; with hairs 9–16 mm; seta
arcuate, 45–75 mm, hairs in lower part of seta 45–55 mm.
Phenology. Stipa 3subdrobovii flowers from May to
June.
Figure 11. Stipa 3subdrobovii M. Nobis & A. Nowak. —A. Holotype. —B. Top of lemma with lower part of awn. —C. Lemma
micromorphological pattern. —D. Callus. —E. Adaxial surface of vegetative leaf in S. 3subdrobovii var. subdrobovii.—F. Adaxial
surface of vegetative leaf in S. 3subdrobovii var. pubescens M. Nobis & A. Nowak. Scale bars: A 55 cm, B, E, F 50.3 mm, C 5
30 mm, D 51 mm.
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Nobis et al. 47
Synopsis of Stipa (Poaceae) in Middle Asia

Habitat. Stipa 3subdrobovii is found in mountain
steppes at 1500–2450 m.
Based on the characteristics of the abaxial surface of
the vegetative leaves (glabrous or pubescent), two va-
rieties of Stipa 3subdodrobovii are recognized.
1. Leaves of vegetative shoots glabrous . . . ........
Stipa 3subdrobovii M. Nobis & A. Nowak var. subdrobovii
19. Leaves of vegetative shoots densely pubescent . . .
Stipa 3subdrobovii var. pubescens M. Nobis & A. Nowak
Stipa 3subdrobovii var. subdrobovii.
Culm leaves: sheaths glabrous or the lower shortly
pubescent, blades glabrous abaxially and pubescent adax-
ially. Leaves of vegetative shoots: ligules up to 0.2 mm,
densely up to 1.5 mm ciliate; abaxial surface of blades
glabrous and smooth or pubescent, whereas adaxial surface
densely covered with hairs 0.15–0.25 mm.
Habitat. Stipa 3subdrobovii var. subdrobovii is
found in high mountain steppes and stony and clay
slopes at 1500–2450 m.
Distribution. Stipa 3subdrobovii var. subdrobovii is
found in Tajikistan (Zeravshan Mountains), western
Kyrgyzstan, and southern Kazakhstan (western Tian-
Shan Mountains).
Additional specimens studied. KAZAKHSTAN. Almatiis-
kai region, 7 km S of Boguto, 5 June 1985, V. Grubov et al. 76
(KRA 455878!, KRA 479063!, KRA 479064!); sandy grass-
land, 15 km N of Kapshagai (40 km N of Almaty), 44°139130N/
77°429170E, 940 m, wp. 266, 22 May 2014, M. Nobis & P.
Gudkova s.n. (KRA!). TAJIKISTAN. Islanderdarya River val-
ley, Zeravshan Region B, western Tajikistan, high mtn. steppe,
among stones on the left slope of the Iskanderdarya River
valley, ca. 0.5 km E of Serimadarun Lake (near Iskanderkul
Lake), 39°059N / 68°239E, 2320 m, incl. SW, slope 5°, 30 May
2015, no. 8, M. Nobis s.n. (KRA 456383!); Iskanderdarya River
valley, Zeravshan Region B, western Tajikistan, Pamir-Alai
Mtns., high mtn. steppe, betw. stones, on the left slope of the
Iskanderdarya River valley, ca. 0.5 km E of Serimadarun Lake
(near Iskanderkul Lake), 39°059080N / 68°229460E, 2326 m,
incl. W, slope 5° to 10°, 14 June 2011, no. 3, M. Nobis s.n.
(KRA 455908!); Zeravshan Mtns., Jagnob River valley, high
mtn. steppe on the right slope of the no name tributary of the
Fagnob River, E of Marzich settlement (W of Anzob),
39°109540N / 68°439490E, 2200–2300 m, exp. W, slope
40°, 22 June 2009, M. Nobis s.n. (KRA!); Fan Mtns., Pamir
Alai Mtns., high mtn. stony steppe, on the mtn. slopes, near N
part of Iskanderkul Lake, 39°059060N / 68°229230E, 2200 m,
inclination S–SSE, 4 June 2009, M. Nobis s.n. (KRA 479069!,
KRA 479062!); Zeravshan River valley, Zeravshan Region C,
Zeravshan Mtns. (Pamiro-Alai); steppe grasslands on the right
terrace of the Zeravshan River, ca. 3–4 km W of Yarm, betw.
Yarm & Langarshif streams (ca. 125 km E of Aini), 39°269100N
/ 69°569040E, 2455 m, exp. S, slope 10° to 60°, no. 4, 20 June
2009, M. Nobis s.n. (KRA 479061!).
Stipa 3subdrobovii var. pubescens M. Nobis & A.
Nowak, var. nov. TYPE: Western Tajikistan, west-
ern Pamir Alai, Zeravshan Mtns., high mtn. steppe,
on the southern slope of mtns. (left slope of the
Iskanderdarya River valley), ca. 0.6 km E of
Serimadarun Lake (near Iskanderkul Lake),
39°059N / 68°239E, 2340 m, 10 June 2012, no.
4/1, M. Nobis & A. Nowak s.n. (holotype, KRA
479068!; isotypes, KRA 407896!, KRA 479065!,
KRA 479066!, KRA 479067!).
Diagnosis. Stipa 3subdrobovii M. Nobis & A. Nowak var.
pubescens M. Nobis & A. Nowak differs from S. 3subdrobovii
var. subdrobovii in having pubescent not glabrous leaves of the
vegetative shoots.
Culm leaves: sheaths glabrous or lower shortly pu-
bescent, blades glabrous abaxially and pubescent adax-
ially. Leaves of vegetative shoots: ligules up to 0.2 mm,
densely ciliate, cilia up to 1.5 mm; abaxial and adaxial
surfaces of blades pubescent and covered with hairs
0.25–0.4 mm.
Habitat. Stipa 3subdrobovii var. pubescens is found
in steppes and stony and clay slopes at 2200–2450 m.
Distribution. In Middle Asia, Stipa 3subdrobovii
var. pubescens is found Tajikistan, within the range of
the nominal variety (Zeravshan Mountains).
Paratypes. TAJIKISTAN. Islanderdarya River valley, Zer-
avshan Region B, western Tajikistan, Pamir-Alai Mtns., high
mtn. steppe, among stones on the left slope of the Iskander-
darya River valley, ca. 0.5 km E of Serimadarun Lake (near
Iskanderkul Lake), 39°059N / 68°239E, 2320 m, inclination
SW, slope 5°, 30 May 2015, no. 8, M. Nobis s.n. (KRA [10
sheets]); western Tajikistan, western Pamir Alai, Zeravshan
Mtns., high mtn. steppe, on the southern slope of mtns. (left
slope of the Iskanderdarya River valley), ca. 0.6 km E of
Serimadarun Lake (near Iskanderkul Lake), 39°059N/
68°239E, 2300 m, inclination SW, slope 5°, 15 June 2012,
no. 35/1, M. Nobis s.n. (KRA 479068, KRA 407893); Islan-
derdarya River valley, Zeravshan Region B, western Tajikistan,
Pamir-Alai Mtns., steppe, among stones on the left slope of the
Iskanderdarya River valley, ca. 0.5 km E of Serimadarun Lake
(near Iskanderkul Lake), 39°059040N / 68°229470E, 2350 m,
17 June 2010, no. 5, M. Nobis s.n. (KRA 479060); steppe
grasslands on the left slope of Iskanderkul Lake, ca. 500 m
NNE of tourist camp Varzob, 39°059560N / 68°209540E,
2250 m, 14 June 2007, no. 2/32, M. Nobis s.n. (KRA
479058, KRA 479059); steppe grasslands on the left slope
of Iskanderkul Lake, ca. 500–1500 m SW of tourist camp
Varzob, 39°059050N / 68°229050E, 2220 m, 12 June 2007, no.
2/32, M. Nobis, M. Kozak & A. Nowak s.n. (KRA 479057).
Notes. Because this variety has densely pubescent
leaves, Stipa drobovii var. iskanderkulica is thought to
be one of its putative parental taxa. Stipa drobovii var.
iskanderkulica also has densely pubescent leaves and
frequently co-occurs with S. subdrobovi var. pubescens at
the same localities. Further examination using molec-
ular methods is needed to evidence the origin of this
taxon.
Stipa subeffusa Ohwi, Acta Phytotax. Geobot. 17: 15.
1957. TYPE: Pakistan. (Hunza) Inter Minapin et
Chalt, 22 Aug. 1955, Nakao s.n. (holotype, KYO
48 Annals of the
Missouri Botanical Garden

not seen; isotype, KYO not seen). 5Achnathe-
rum brandisii (Mez) Z. L. Wu, Acta Phytotax. Sin.
34: 154. 1996 [syn. after Freitag, 1985].
Stipa subsessiliflora (Rupr.) Roshev., Izv. Imp. Bot.
Sada Petra Velikago 14(Suppl. 2): 50. 1915.
Basionym: Lasiagrostis subsessiliflora Rupr., Sert.
Tianschan. publ. in M ´em. Acad. Imp., St. P ´etersb.,
S´er. 7, 14(4): 35. 1869. TYPE: [China. Xinjiang:]
Mittlerer Tianshan, Toyandy-Tal (ca. 70 km NNW
of Kashgar), 30 July 1886, Osten-Sacken s.n. (ho-
lotype, LE!).
Synonyms. [Ptilagrostis subsessiliflora (Rupr.)
Roshev., Fl. URSS 2: 74. 1934.
Habitat. Stipa subsessiliflora is found in high moun-
tain steppes and stony slopes at 3000–4200 m.
Distribution. Stipa subsessiliflora is found in the
mountains of central Asia (Tzvelev, 1968, 1976; Wu
& Phillips, 2006). In Middle Asia, it occurs in north-
eastern Afghanistan, western China, Kyrgyzstan, east-
ern Tajikistan, and eastern Pakistan.
Stipa szovitsiana Trin. in Hohenacker, Bull. Soc. Imp.
Naturalistes Moscou 11(3): 243. 1838. TYPE:
Tatuni, July 1853, “mis. cl.,”Hohenacker 1253
(lectotype, designated by Tzvelev, [1976: 584],
LE!). 5Stipa arabica Trin. & Rupr. var.
arabica.
Stipa 3tadzhikistanica M. Nobis, Pl. Syst. Evol.
299(7): 1338. 2013 (S. caucasica Schmalh. 3S.
lipskyi Roshev.; Nobis, 2013). TYPE: Dolina rzeki
Iskanderdaria (Iskanderdarya River valley), Rejon
Zerawsza´nski B (Zeravshan Region B), Zachodni
Tad˙zykistan (West Tajikistan) Pamiro-Alai, mur-
awy stepowe w´sr´od głaz ´ow, na stokach g ´or, na
lewym zboczu doliny rzeki Iskanderdarya, ok.
0.5 km na E od jeziora Serimadarun (koło Jeziora
Iskanderkul [Tajikistan, Zeravshan Mtns. (Pamir
Alai Mtns.), steppe grasslands on the mtn. slopes,
among stones, on the left slope of the Iskander-
darya River valley, ca. 0.5 km E of the Serima-
darun Lake (near Iskanderkul Lake)], 39°059040N
/ 68°229470E, 2350 m, exp. SE, slope 10°, no. 5,
17 June 2010, M. Nobis s.n. (holotype, KRA
383654!; isotypes, Herb. Stip. M. Nobis!, KRA!,
KRAM!).
Habitat. Stipa 3tadzhikistanica is found in steppe
grasslands at 1400–2400 m.
Distribution. Stipa 3tadzhikistanica is found in
Middle Asia (Nobis, 2013) in Tajikistan (Hissar and
Zeravshan Mountains).
Stipa 3talassica Pazij, Bot. Mater. Gerb. Inst. Bot.
Zool. Akad. Nauk Uzbeksk. S.S.R. [Not. Syst.
Herb. Inst. Bot. & Zool. Acad. Sci. Uzbek.] 10:
21. 1948 (S. caucasica Schmalh. 3S. macroglossa
P. A. Smirn. subsp. macroglossa; Nobis, 2013).
TYPE: Tsentralnyi Tyan-Shan, Verkhne-Talasskii
raion, Talasskii Alatau, Dol. r. Besh-tash, Poyas
khvoinogo lesa, skaly, 30 June 1927, No. 339, M.
M. Sovetkina & M. V. Uspenskaya s.n. (holotype,
TASH 163462!; isotype, LE!).
Habitat. Stipa 3talassica is found in steppe grass-
lands at 1100–1800 m.
Distribution. Stipa 3talassica is found in Middle
Asia (Pazij, 1968; Nobis, 2013) in Kyrgyzstan (Talass,
Kyrgyz, and Fergana Mountains).
Stipa tianschanica Roshev., Fl. Aziat. Ross. 12: 149,
pl. 10, f. 3, 3a. 1916. TYPE: Semirech. obl.
Przhev. u. r. Ak“-Shiiryak”nizove, kamenistye
sklony, 31 July 1913, B. Shishkin s.n. (lectotype,
designated by Nobis [2014: 302], MW!).
Habitat. Stipa tianschanica is found in high moun-
tain steppes, semideserts, and rocky ledges and crevices
at 2500–4000 m.
Distribution. Stipa tianschanica is found in the
mountains of central Asia (Nobis, 2011, 2014). In Middle
Asia, it occurs in western China (Tian-Shan, Pamir,
Karakorum Mountains), Kyrgyzstan (Tian-Shan Moun-
tains), and Tajikistan (Pamir Mountains). Figure 4.
Notes. Within the species, the two subspecies can be
distinguished: Stipa tianschanica subsp. tianschanica
(which occurs in the studied area) and S. tianschanica
subsp. gobica (Roshev.) D. F. Cui (synonyms: [S. gobica
Roshev., [S. tianschanica var. gobica (Roshev.)P.C.Kuo
& Y. H. Sun). Since the latter taxon grows in northeastern
Kazakhstan (Altay Mountains) and western China (Nobis,
2014) near the borders of the studied area, it is also
possible to find it in Middle Asia. Stipa tianschanica
subsp. tianschanica differs from S. tianschanica subsp.
gobica in having glumes 24–34 mm long (vs. 14–27 mm
long), the top of the lemma with a well-developed ring of
hairs (vs. the top of the lemma without a ring of hairs or,
rarely, with a few scattered, short hairs), awn (63–)
75–90(2103) mm long (vs. awn [42–]55–80 mm long)
and the seta (51–)60–80(286) mm long (vs. seta [31–]
40–60[270] mm long).
Stipa tibetica Mez, Repert. Spec. Nov. Regni Veg. 17:
207. 1921. TYPE: Lasiag. Mongholica Trin., Tibet
Occ., alt. 14,000 ft., T. T. (Herb. Ind. Or. Hook. fil.
& Thomson) (lectotype, designated here,
LE00009272!; holotype, B†). [Ptilagrostis
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Nobis et al. 49
Synopsis of Stipa (Poaceae) in Middle Asia

tibetica (Mez) Tzvelev, Rast. Tsentr. Azii 4: 45.
1968.
Stipa tirsa Steven, Bull. Soc. Imp. Naturalistes Moscou
30(2): 115. 1857. TYPE: Ukraine. Kaltschik, camp.
Maeotic, Graff s.n. (lectotype, designated by
Martinovsk´y & Skalick ´y [1969: 339], H not seen).
Synonyms. [Stipa pennata L. var. tirsa (Steven)
ˇ
Celak., Sitzungsber. K¨onigl. B¨ohm. Ges. Wiss. Prag,
Math.-Naturwiss. Cl. 1884: 58. 1884.
Notes. Although we did not find any specimens of
Stipa tirsa during this study, it is known from northern
Kazakhstan (Tzvelev, 1976), so it is also very likely to be
found in the area of study. Narrow-leaved specimens of
S. zalesskii sometimes may be confused with S. tirsa.
However, S. tirsa has vegetative leaves with a bristle-
like apex and very short (0.1–0.2 mm long) ligules,
whereas in S. zalesskii vegetative leaves are obtuse at the
apex and ligules are always longer.
Stipa tortilis Desf., Fl. Atlant. 1: 99–100, pl. 31, f. 1.
1798. TYPE: in arvis (holotype, P not seen, syn.
after Freitag, 1985). 5Stipellula capensis
(Thunb.) R¨oser & Hamasha, Schlechtendalia 24:
92. 2012.
Stipa transcaucasica Grossh., Trudy Bot. Inst. (Baku) 2:
245–246. 1936. TYPE: [Azerbaijan.] Zuvant, Kis-
Jurdy Mtns. near Gevedara, 17 July 1930, Prilipko
s.n. (holotype, BAK not seen). [Stipa holoser-
icea Trin. var. transcaucasica (Grossh.) M.
Nobis.
Stipa trichoides P. A. Smirn., Repert. Spec. Nov.
Regni Veg. 21: 233. 1925. TYPE: Turkomania.
Pr. Ashabad. In m. Ludsha, 6500, 9 July 1898, D.
Litwinow 2222 (lectotype, designated by Nobis
et al. [2016a: 150], LE!; isolectotypes, LE! [3
sheets], MW!).
Synonyms. [Stipa turkestanica Hack. subsp. tri-
choides (P. A. Smirn.) Tzvelev, Novosti Syst. Vyssh.
Rast. 11: 17. 1974.
Habitat. Stipa trichoides is found in steppes, stony
slopes, and screes at 1800–2900 m.
Distribution. Stipa trichoides is found in Middle and
southwestern Asia (Gonzalo et al., 2013; Nobis et al.,
2016a). In Middle Asia, it occurs in Afghanistan, north-
ern Iran, Kyrgyzstan, Tajikistan, Turkmenistan, and
Uzbekistan.
Stipa turcomanica P. A. Smirn., Repert. Spec. Nov.
Regni Veg. 21: 234. 1925. TYPE: Pl. Turkoma-
niae, in montibus prope Ashabad, 1000 m, 9 May
1897, D. Litvinov 177 (lectotype, designated by
Tzvelev [1976: 588], LE!; isolectotypes, LE!). [
Stipa zalesskii Wilensky var. turcomanica
(P. A. Smirn.) M. Nobis.
Stipa turgaica Roshev., Bot. Mater. Gerb. Bot. Inst.
Komarova Akad. Nauk S.S.S.R. 11: 21–22. 1949.
TYPE: Turgai Distr. & post, Kizyl-Jingilskaya
Volost, series of small lakes in the lower reaches
of Sarysu, Lake Tailyak-Kul, wormwood-biyurgun
steppe, 21 May 1914, I. Krasheninnikov 5104
(lectotype, designated by Tzvelev [1976: 584],
LE!). [Stipa arabica Trin. & Rupr. var. turga-
ica (Roshev.) Tzvelev.
Stipa turkestanica Hack., Trudy Imp. S.-Peterburgsk.
Bot. Sada. 26: 59. 1906. TYPE: Iter Turkestani-
cum 1904; Stipa turkestanica Hack., Shugnan:
stoyanka Dzhidaka, 27 July 1904, B. A. Fedt-
schenko s.n. (holotype, W-Hackel No. 19184!;
isotypes, MW! [2 sheets]).
Habitat. Stipa turkestanica is found in steppes,
stony slopes, and screes at 1800–4000 m.
Distribution. Stipa turkestanica is found in the
mountains of central and southwestern Asia (Nobis
et al., 2016a). In Middle Asia, it occurs in Afghanistan,
western China, northern Iran, eastern Pakistan, and
Tajikistan. Figure 3.
Notes. Stipa turkestanica var. diyaensis L. Q. Zhao
& K. Guo has recently been described from western
China (Himalayas). It differs from the nominal variety
only in the character of its awn column, which is shortly
pubescent versus glabrous. During our herbarium visits,
we found another specimen of this taxon, collected from
southeastern Afghanistan (Prov. Paktia, 3 km S of Zirok,
rd. from Khost to Urgun, 29 May 1971, O. H. Volk 71/
188 [M-0257951], rev. M. Nobis). Despite S. turkestan-
ica var. diyaensis being unknown from the research area
up to this point, its occurrence in Middle Asia is very
likely.
Stipa 3tzveleviana Kotukhov, Bot. Zhurn. (Moscow &
Leningrad) 79(7): 102. 1994 (S.orientalis Trin. 3
S.macroglossa P. A. Smirn. subsp. kazachstanica
(Kotukhov) M. Nobis; Nobis & Gudkova, 2016).
TYPE: Saur-Tarbagatai, brachia australi-
occidentalia jugi Manrak, 600 m, locus Kempir-
bulak, clivulus saxosus boreali-occidentalis, 11
July 1992, Ju. Kotuchov s.n. (holotype, LE!; iso-
types, LE! [3 sheets]).
Habitat. Stipa 3tzveleviana is found in steppe
grasslands at 600–1400 m.
Distribution. Stipa 3tzveleviana is found in the
mountains of northeastern Middle Asia (Kotukhov,
50 Annals of the
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2002; Nobis & Gudkova, 2016). It occurs specifically in
eastern Kazakhstan (Sauro-Manrak Mountains).
Stipa 3tzvelevii Ikonn., Opred. Vyssh. Rast. Badakh-
shana 84. 1979 (S. caucasica Schmalh. 3S.
orientalis Trin.; Nobis, 2011). TYPE: Tajikistan.
Badakhshan, Kishty Dzharob Valley, near Lyangar
settlement, NE exposure, 3700 m, 10 July 1956, G.
Ladygina 3219 (holotype, LE!).
Habitat. Stipa 3tzvelevii is found in steppe grass-
lands, screes, rocks, and stony slopes at 1500–2800 m.
Distribution. Stipa 3tzvelevii is found in Middle
Asia (Nobis, 2011) in Kazakhstan, Kyrgyzstan, and
Tajikistan.
Stipa ucrainica P. A. Smirn., Repert. Spec. Nov. Regni
Veg. 22: 374. 1926. TYPE: Ekaterinoslavskaya
gub., Aleksandrovskii u., bliz khut. Mirgorodovka,
stepnoi sklon Ternovoi balki, V. Alekhin 209 (ho-
lotype, MW!; isotypes, MW! [4 sheets]).
Synonyms. [Stipa zalesskii Wilensky subsp.
ucrainica (P. A. Smirn.) Tzvelev, Novosti Sist. Vyssh.
Rast. 11: 17. 1974; 5S. krascheninnikowii Roshev.
Habitat. Stipa ucrainica is found in steppes and
stony slopes at 300–1000 m.
Distribution. Stipa ucrainica is a widespread spe-
cies. Its range extends from eastern Europe to the
northern part of Middle Asia (Tzvelev, 1976; Slyusar-
enko, 1977). In Middle Asia it occurs in western and
central Kazakhstan.
Stipa ukranensis Lam., Tabl. Encycl. 1: 157. 1791.
TYPE: Ex Ukrania, Tirsa, Guettard mem. v. 1. t.
1.2 (not seen; syn. after Tzvelev, 1976). 5Stipa
capillata L.
Stipa violacea Nikitina, Trudy Biol. Inst. Kirg. Fil. An
SSSR 2: 68. 1947, hom. illeg. non Hitchcock,
Contr. U.S. Natl. Herb. 24(7): 282. 1925. TYPE:
Tsentralnyi Tian-shan, severnyi sklon khr. Kavak-
tau, pereval mezhdu Beiryuk i Kash-bel, 28 July
1937, E. Mikhailova & L. Popova 66 (holotype,
LE!). 5Stipa kirghisorum P. A. Smirn.
Stipa voronini Krasn., Spisok. Rast. Sobr. Vost. Tyan-
Shane, letom 1886 goda 125, 1887. TYPE: Fl.
Iliensis, prope Andraki, May 1886, Krassnov s.n.
(lectotype, designated by Tzvelev [1976: 578],
LE!). 5Stipa richteriana Kar. & Kir. subsp.
richteriana.
Stipa woroninii Krasn., Script. Hort. Univ. Petrop. II, 1:
22. 1887–1888. TYPE: Fl. Iliensis, prope
Andraki, May 1886, Krassnov s.n. (lectotype,
designated by Tzvelev [1976: 578], LE!). 5Stipa
richteriana Kar. & Kir. subsp. richteriana.
Stipa zalesskii Wilensky, Dnevn. Vserossisk. S”ezda
Russk. Bot. 41. 1921. TYPE: Kazakhstan. In the
vic. of Saratovka, southern slopes in the regions of
Kalyubanov’s country house, 5 June 1918, D.
Zalesskii s.n. (lectotype, designated by Tzvelev
[1976: 587], LE!).
Within the species three varieties are recognized.
1. Awn glabrous or scabrous on column and plumose
on seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
19. Awn pilosethroughout.........................
..... Stipa zalesskii Wilensky var. iljinii (Roshev) Tzvelev
2. External sheaths of vegetative shoots shortly pubes-
cent . . ..................Stipa zalesskii var. zalesskii
29. External sheaths of vegetative shoots glabrous . . .
Stipa zalesskii var. turcomanica (P. A. Smirn.) M. Nobis
Stipa zalesskii var. zalesskii.
Synonyms. [Stipa pennata L. subsp. zalesskii
(Wilensky) Freitag, Notes Roy. Bot. Gard. Edinburgh
42: 443. 1985; 5S. rubens P. A. Smirn.; [S. zalesskii
Wilensky var. rubens (P. A. Smirn.) Tzvelev, Novosti Sist.
Vyssh. Rast. 11: 19. 1974; 5S. rubentiformis P. A.
Smirn.; [S. rubens P. A. Smirn. subsp. rubentiformis (P.
A. Smirn.) F. M. V ´azquez & M. Gut., Telopea 13(1–2):
164. 2011; 5S. rubens subsp. sublevis Martinovsk´y,
Preslia 44(1): 21. 1972. TYPE: Krymskaja oblast, Sirn-
feropolskyj ra-n, s. Partizanske, l July l955, A. Barbaricz,
D. Dobroczyaeva & M. Kukalo s.n. (holotype, KWHA not
seen); 5S. maeotica Klokov & Osychnyuk; [S. zalesskii
var. maeotica (Klokov) Tzvelev, Byull. Moskovsk.
Obshch. Isp. Prir., Otd. Biol. n.s. 91(1): 121. 1986; 5
S. smirnovii Martinovsk´y; 5S. canescens P. A. Smirn. ex
Roshev.
Habitat. Stipa zalesskii var. zalesskii is found in
steppe grasslands at 1300–3200 m.
Distribution. Stipa zalesskii var. zalesskii is a widely
distributed species. Its range extends from central
Europe up to central Asia (Tzvelev, 1976; Conert,
1998; Nobis et al., 2016b, 2016c). In Middle Asia,
it occurs in western China, Kazakhstan, and
Kyrgyzstan.
Stipa zalesskii var. iljinii (Roshev.) Tzvelev, Novosti
Sist. Vyssh. Rast. 11: 18. 1974. Basionym: Stipa
iljinii Roshev., Bull. Jard. Bot. Acad. Sc. URSS
1931, 30: 294. 1932.
Habitat. Stipa zalesskii var. iljinii is found in steppe
grasslands, within the altitudinal range of the nominal
variety.
Distribution. Stipa zalesskii var. iljinii is found
within the range of the nominal variety (Roshevitz,
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Nobis et al. 51
Synopsis of Stipa (Poaceae) in Middle Asia

1916; Tzvelev, 1976). In Middle Asia it occurs in
Kazakhstan.
Notes. See comments for Stipa borysthenica var.
anomala.
Stipa zalesskii var. turcomanica (P. A. Smirn.) M.
Nobis, comb. & stat. nov. Basionym: Stipa turco-
manica P. A. Smirn., Repert. Spec. Nov. Regni
Veg. 21: 234. 1925.
Synonyms. [Stipa zalesskii subsp. turcomanica
(P. A. Smirn.) Tzvelev, Nov. Syst. Vyssh. Rast. 11:
18. 1974.
Habitat. Stipa zalesskii var. turcomanica is found in
steppe grasslands at 1200–2000 m.
Distribution. Stipa zalesskii var. turcomanica is
found in the southern part of Middle Asia (Tzvelev,
1976) in northern Iran and Turkmenistan (Kopet-Dagh
Mountains).
Notes. The only difference between Stipa zalesskii
var. turcomanica and S.zalesskii var. zalesskii is the
character of the sheaths of the vegetative shoots, which
are glabrous versus shortly pubescent, respectively.
This character (pubescence or lack of indumentum)
is often considered natural variation, and both glabrous
as well as pubescent sheaths can be observed within one
population, e.g., in S. caucasica. However, we did not
notice such variation in specimens identified as S.
zalesskii var. zalesskii or S. zalesskii var. turcomanica,
whereas the length of the awn and anthecium,
considered a diagnostic character by Tzvelev (1976),
varies considerably across the range of these taxa (Nobis
at al., 2016b, 2016c; M. Nobis, pers. obs.).
Stipa zeravshanica M. Nobis, Nordic J. Bot. 31(6):
667. 2013. TYPE: Western Tajikistan, western
Pamir Alai Mtns.: G´ory Zerawsza ´nskie, w szczeli-
nach i na p´ołkach skalnych, na prawym, skalistym
stoku doliny rzeki Jagnob (przy drodze), ok. 5 km
na W od wsi Anzob (pomie
˛dzy osadami Anzob i
Marzicz) [Zeravshan Mtns., in rocky crevices and
ledges, on the right slope of the Jagnob River valley
(near the rd.), ca. 5 km W of Anzob (betw. Anzob &
Marzich settlements)]; 39°109490N / 68°469490E;
2100 m, incl. S, slope 80°, 10 June 2009, M. Nobis
3a (holotype, KRA 407887!; isotypes, Herb. Stip.
M. Nobis! [2 sheets], KRA 407870!, KRA
407871!, KRA 407884!, KRA 407885!, KRA
407886!, KRA 407858!, KRAM!, WA!).
Habitat. Stipa zeravshanica is found in ledges and
fissures of calcareous rocks at 1500–2600 m.
Distribution. Stipa zeravshanica is found in Middle
Asia (Nobis et al., 2013), specifically in Tajikistan
(Zeravshan Mountains, northern part of Hissar Moun-
tains, and northern Turkestan Mountains). Figure 3.
TAXONOMIC-PHYTOGEOGRAPHIC ANALYSIS OF THE MIDDLE
ASIAN FEATHER GRASSES
In Middle Asia the genus Stipa comprises 98 taxa,
including 72 species, four subspecies, and 22 varieties.
Of the 72 species of feather grasses, 23 are of hybrid
origin (nothospecies). For all nothospecies we indicated
putative parental species. Several other taxa (e.g., S.
kempirica,S. korshinskyi,S. kotuchovii,S. okmirii,S.
sczerbakovii) are probably of hybrid origin as well, but
further studies using molecular methods are needed to
establish this. The number of feather grasses occurring
in different countries varies considerably. The countries
richest in Stipa taxa (species and subspecies) include
Kazakhstan (42 taxa), Tajikistan (40 taxa), and Kyrgyz-
stan (35 taxa; Table 2, Fig. 12).
Among the 76 taxa (species and subspecies) of Stipa
recorded in Middle Asia, 41 are endemic, with distribu-
tion restricted either to a particular country or mountain
range or to the region in general. Eight species (S.
himalaica,S. kirghisorum,S. korshinskyi,S. lingua,S.
roborowskyi,S. tianschanica,S. trichoides,andS. turkes-
tanica) distributed in Middle Asia, with several stations
exceeding the borders of the studied region, can be
considered subendemics. Among endemics recorded in
the research area, 20 taxa are of hybrid origin (Table 2).
The highest number of endemic taxa of Stipa has been
recorded in southern Kazakhstan (21 taxa: 12 species,
nine nothospecies), Tajikistan (20 taxa: 11 species, nine
nothospecies), Kyrgyzstan (14 taxa: 10 species, four
nothospecies), and Uzbekistan (seven taxa: six species,
one nothospecies).
Based on sectional divisions of the genus Stipa pro-
posed by Tzvelev (1974), Middle Asian feather grasses
can be divided into six sections (Fig. 13): Smirnovia
Tzvelev (13 species, one subspecies, 12 nothospecies),
Barbatae A. Junge (11 species, four nothospecies), Stipa
(11 species, two subspecies, two nothospecies), Leiostipa
Dumort. (10 species, one subspecies, five nothospecies),
Pseudoptilagrostis Tzvelev (three species), and Regelia
Tzvelev (one species). However, in accordance with the
results of our numerical analyses, sectional affiliations of
some taxa, especially those representing sectionBarbatae
(e.g., S. purpurea,S. orientalis,S. gracilis,S. himalaica,
S. zeravshanica,S. 3gnezdilloi,S. breviflora,S. robor-
owski) as well as those from section Leiostipa
(S. 3dzungarica,S. 3heptapotamica,S. breviflora,
S. richteriana; Fig. 13) are puzzling. Some of the species
mentioned previously in this article create subclades that
are clearly separated from the other ones (Freitag, 1985).
Because of the hybrid origin of some taxa studied, their
sectional affiliations are problematic, and sometimes
52 Annals of the
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Table 2. List of Stipa L. taxa (species and subspecies) occurring in Middle Asian countries. The presence of a taxon in a
particular country is marked using a plus sign (1), endemics are listed in bold, and subendemics are marked with an asterisk (*).
N
Afghanistan
W
China
N
Iran Kazakhstan Kyrgyzstan
N
Pakistan Tajikistan Turkmenistan Uzbekistan
S. 3adamii ...1... . .
S. aktauensis .... . . . . 1
S. 3alaica 1.. . 1.11 .
S. 3albasiensis .1.11.1..
S. arabica 1111 1 1 1 1 1
S. 3assyriaca 1.11 1 .11 1
S. badachschanica 1.1...1..
S. 3balkanabatica .... . . . 1.
S. basiplumosa .1.. . 1.. .
S. borysthenica .1.1... . .
S. 3brevicallosa .... . .1..
S. breviflora .1.. 11.. .
S. 3brozhiana .... . .1..
S. bungeana 11.. 11.. .
S. capillata 1111 1 1 1 1 1
S. caucasica subsp.
caucasica
1111 1 .11 1
S. caucasica subsp.
nikolai
1.. 11.1.1
S. 3consanguinea .1.1... . .
S. 3czerepanovii ...1... . .
S. dasyphylla ...1... . .
S. drobovii 1.11 1 .11 1
S. 3dzungarica ...1... . .
S. 3fallax .... . .1..
S. glareosa 11.1111 ..
S. 3gnezdilloi .... . . . . 1
S. gracilis .1.11.1..
S. 3heptapotamica ...1... . .
S. himalaica* 11.. . 1.. .
S. 3hissarica .... . .1..
S. hohenackeriana 1111 1 1 1 1 1
S. holosericea ..1....1.
S. 3kamelinii ...1... . .
S. karakabinica ...1... . .
S. karataviensis ...1... . 1
S. kempirica ...1... . .
S. kirghisorum* 11.1111 .1
S. korshinskyi* ...1... . .
S. kotuchovii ...1... . .
S. krylovii .1.11.1..
S. lessingiana .11 1 1 .11 1
S. lingua* 1.1...11 1
S. lipskyi ...11.1.1
S. longiplumosa .... 1.1.1
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Table 2. Continued.
N
Afghanistan
W
China
N
Iran Kazakhstan Kyrgyzstan
N
Pakistan Tajikistan Turkmenistan Uzbekistan
S. macroglossa
subsp.
kazachstanica
.1.11.. . .
S. macroglossa
subsp.
macroglossa
...11.1.1
S. magnifica .... 1.. . .
S. 3manrakica ...11.. . .
S. margelanica 1.. . 1.1..
S. narynica .... 1.. . .
S. okmirii .... . .1..
S. orientalis 1111 1 1 1 .1
S. ovczinnikovii .... . .1..
S. penicillata .1.. . . . . .
S. 3pseudo-
capillata
...1... . .
S. 3pseudo-
macroglossa
.... . .1..
S. pulcherrima
subsp. crassiculmis
..1....1.
S. pulcherrima subsp.
pulcherrima
...11.. . .
S. purpurea 11.. 111 ..
S. regeliana 11.1111 ..
S. richteriana
subsp. jagnobica
1.. . . . 1..
S. richteriana
subsp.
richteriana
.1.1... . 1
S. roborowskyi* .1.. . 1.. .
S. sareptana .1.11.11 1
S. sczerbakovii ...1... . .
S. 3subdrobovii ...1..1..
S. subsessiliflora 11.. 111 ..
S. 3tadzhikistanica .... . .1..
S. 3talassica .... 1.. . .
S. tianschanica* .1.. 1.1..
S. trichoides* 1.1.1.11 1
S. turkestanica* 111..11 ..
S. 3tzveleviana ...1... . .
S. 3tzvelevii ...11.1..
S. ucrainica ...1... . .
S. zalesskii .11 1 1 .. 1.
S. zeravshanica .... . .1..
Total no. of taxa 22 28 15 42 35 15 40 15 20
Endemics 5 3 1 21 13 0 20 2 7
54 Annals of the
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these taxa have been incorporated into one or another
section. For instance, most taxa originating from the
hybridization of species representing sections Stipa
and Smirnovia (e.g., S. 3manrakica,S. 3alaica,S.
3talassica or S. 3pseudomacroglossa), as well as sec-
tions Leiostipa and Smirnovia (S. 3consanguinea and
Figure 12. Number of Stipa L. taxa in particular countries of Middle Asia; black columns refer to species and subspecies, gray
refers to nothospecies.
Figure 13. —A, B. Cluster analysis (UPGMA method of classification and Gower’s general similarity coefficient) performed on
16 qualitative characters. As an outgroup to Stipa L., selected members from the genera Achnatherum P. Beauv., Ptilagrostis
Griseb., Neotrinia (Tzvelev) M. Nobis, P. D. Gudkova & A. Nowak, Orthoraphium Nees, Trikeraia Bor, and Piptatherum P. Beauv.
were used. Lemma epidermal patterns are presented for the following species: (a) Achnatherum mandaivillei (Freitag) M. Nobis
(Oman, Mandaville 6525 [KAS]), (b) Ptilagrostis concinna (Hook. f.) Roshev. (India, Ladakh, L. Klimeˇs s.n. [KRA]), (c) Neotrinia
splendens (Trin.) M. Nobis, P. D. Gudkova & A. Nowak (Tajikistan, Yu. Gusev s.n. [LE]), (d) Orthoraphium roylei Nees (Nepal, M. A.
Farille s.n. [E]), (e) Trikeraia hookeri (Stapf) Bor (China, Tibet [PE 718306]), (f) Piptatherum microcarpum (Pilg.) Tzvelev
(Tajikistan, M. Nobis [KRA]), (g) Stipa ucrainica P. A. Smirn. (Kazakhstan, R. Roshevitz et al. s.n. [LE]), (h) S. lessingiana Trin. &
Rupr. (Kazakhstan, M. Nobis s.n. [KRA]), (i) S. karataviensis Roshev. (Kazakhstan, R. Kamelin et al. s.n. [LE]), (j) S. richteriana
Kar. & Kir. (Kazakhstan, M. Nobis s.n. [KRA]), (k) S. korshinskyi Roshev. (Kazakhstan, V. Goloskokov s.n. [LE]), (l) S.
badachschanica Roshev. var. pamirica (Roshev.) M. Nobis (Tajikistan, M. Krivonogova 256 [TAD]), (m) S. regeliana Hack.
(China, A. A. Yunatov & Yuan U-fen 623 [LE]).
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Synopsis of Stipa (Poaceae) in Middle Asia

S. 3balkanabatica), due to the presence of unigenicu-
late or indistinctly bigeniculate awns, have been in-
cluded by different authors into one or another
sections. This problem also applies to taxa originat-
ing from hybridization between species belonging to
sections Leiostipa and Stipa (e.g., S. 3pseudocapillata
or S. 3heptapotamica) that were usually included in
section Barbatae. Several years ago, Tzvelev (2012, 2014)
distinguished additional sections, Hemibarbatae Tzvelev,
Subbarbatae Tzvelev, and Hybridogenae Tzvelev. Section
Figure 13. (Continued)
56 Annals of the
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Subbarbatae is monotypic and comprises only S. lessingi-
ana, whereas section Hybridogenae comprises taxa orig-
inating from hybridization between species from sections
Stipa and Subbarbatae. Keeping in mind that intersectional
hybrids in Stipa are common and occur between species
from almost all sections distinguished to date, following
Tzvelev’s practice, five or six additional new sections
dedicated only for that nothospecies could be distinguished
Figure 14. Altitudinal distribution of Stipa L. species in Middle Asia.
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Nobis et al. 57
Synopsis of Stipa (Poaceae) in Middle Asia

within the genus Stipa.Eveninthecaseofsome“good”
(pure) species, those for which a hybrid origin has not been
suggested, the sectional affiliation might also be confusing.
For example, S. himalaica,S. zeravshanica,andS. ori-
entalis, due to their morphological characters, i.e., the
presence of an ovary with two styles and pilose awns
throughout, might be included either in section Bar-
batae or section Stipa (cf. Tzvelev, 1974; Freitag,
1985). A similar problem involves several species from
section Hemibarbatae (Tzvelev, 2012) that have been
included previously either in sections Leiostipa or
Barbatae (Tzvelev, 1974; Freitag, 1985).
Today, molecular techniques are widely used for
identifying phylogenetic relations between and within
different genera of Stipeae (e.g., Romaschenko et al.,
2008, 2010, 2012; Hamasha et al., 2012; Sclovich et al.,
2015; Krawczyk et al., 2017, 2018). In accordance with
the preliminary results of phylogenetic studies on the
genus Stipa conducted by Krawczyk et al. (2017, 2018),
S. orientalis is more closely related to S. glareosa and S.
tianschanica, the species belonging to section Smirno-
via, rather than to representatives from sections Barba-
teae or Stipa, with which the taxon was traditionally
classified (Tzvelev, 1976). Another example is S. rich-
teriana s.l., which creates a clade with S. lessingiana
rather than with other representatives of Stipa,Leiostipa,
or Barbatae (Krawczyk et al., 2017), whereas based on
the results of Hamasha et al. (2012) and Krawczyk et al.
(2017), S. pururea and S. roborowskyi are closely related
to species representing section Pseudoptilagrostis, not
section Barbatae, which partially corresponds with the
results of our analysis presented in this paper (Fig. 13).
To resolve the problem of the phylogenetic relationships
between and within particular sections in the entire
genus Stipa, further studies involving the use of molec-
ular methods are needed, and this is the topic of our next
paper (Nobis et al., in prep.).
In Middle Asia, members of the genus Stipa can be
found at elevations from (0 to)300 to 4500(to 5000) m
(Fig. 14), but most of the studied feather grasses are
mountain species. Greatest species richness is observed
at altitudes between 900 and 2500 m, with 31 to 38
species (34 species on average) per every 100 m (Figs.
14, 15). Areas situated above 3000 m are occupied by
19 species, but in the highest elevations, above 4000 m,
only nine species of feather grasses have been found.
Some species have a very narrow altitudinal range, but
others such as S. bungeana,S. glareosa,S. gracilis,S.
hohenackeriana,S. kirghisorum,S. krylovii, and S.
orientalis have a very wide one (Fig. 14). All of these
taxa can be found in numerous localities in the moun-
tains as well as in some of the lowlands, e.g., in central
Kazakhstan (S. glareosa,S. orientalis,S. kirghisorum).
Like other mountain taxa, their current range may be a
result of glaciations and previous climate changes. In
the lowest elevations, up to 900 m, four species occur,
i.e., S. aktauensis,S. kempirica,S. 3pseudocapillata,
and S. borysthenica (Fig. 14). These species grow in
lowlands and in lower mountain ranges within steppes or
deserts in central Uzbekistan and/or central Kazakhstan
(Pazij, 1968; Tzvelev, 1976; Kotukhov, 2002; Nobis,
2012). The majority of feather grasses are typical steppe
taxa, occurring in different lowland and mountain grass-
lands on sand, loess, gravel, and rock. Some of the
analyzed taxa can occupy a few different habitats, e.g.,
steppes, semideserts, dry grasslands, rocky grasslands,
and screes (e.g., S. caucasica,S. drobovii,S. kirghiso-
rum,orS. orientalis). However, there is also a group of
species restricted to one or two habitat types, e.g., in
high mountain deserts (e.g., S. orientalis,S. glareosa,S.
basiplumosa,S. subsessiliflora), sands (e.g., S. arabica,
S. borysthenica), screes (e.g., S. macroglossa), or alpine
meadows (e.g., S. regeliana,S. penicillata,S. purpurea).
Some have a very narrow ecological range, growing only
Figure 15. Altitudinal distribution of feather grasses in Middle Asia (number of species occurring in 100-m altitudinal belts).
58 Annals of the
Missouri Botanical Garden

on rocky habitats (ledges and fissures) on calcareous
substrates (e.g., S. himalaica,S. gracilis,S. zeravshan-
ica). Two of the last mentioned species, namely S.
gracilis and S. zeravshanica, represent typical geograph-
ical vicariants, occurring in isolated mountain ranges.
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Volume 105, Number 1
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Synopsis of Stipa (Poaceae) in Middle Asia
ISSN 0026-6493 (PRINT); ISSN 2162-4372 (ONLINE)