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Webbia
Journal of Plant Taxonomy and Geography
ISSN: 0083-7792 (Print) 2169-4060 (Online) Journal homepage: https://www.tandfonline.com/loi/tweb20
Computer-generated key and descriptions of
Phleum species (Poaceae)
A. El-Gazzar, A. M. Eisa & A. A. Khattab
To cite this article: A. El-Gazzar, A. M. Eisa & A. A. Khattab (2016) Computer-
generated key and descriptions of Phleum species (Poaceae), Webbia, 71:1, 25-35, DOI:
10.1080/00837792.2015.1119955
To link to this article: https://doi.org/10.1080/00837792.2015.1119955
Published online: 13 Jan 2016.
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Computer-generated key and descriptions of Phleum species (Poaceae)
A. El-Gazzar
a
*, A. M. Eisa
b
and A. A. Khattab
c
a
Botany Department, Faculty of Science, Suez Canal University, El-Arish, N. Sinai, Egypt;
b
Botany Department, Faculty of Science,
Damanhour University, El-Beheira, Egypt;
c
Faculty of Science, The Herbarium, Botany Department, Cairo University, Giza, Egypt
(Received 3 September 2015; final version received 12 November 2015)
Conflicting and often cursory taxonomic accounts of the genus Phleum L. (Poaceae: Pooideae) have led to difficulties in
discriminating among the taxa. Apart from the two notho-species Phleum ×brueggeri and Phleum ×viniklarii without
any synonyms, more than 250 synonyms are on record for the remaining 16 species and their infra-specific taxa. The
limited range of characters used in traditional keys to these taxa greatly reduced their applicability. To overcome these
difficulties, the key-generating program suite DELTA was used to construct a conventional key to 15 Phleum species (of
which five are represented by two subspecies each), based on 20 gross morphological and anatomical characters system-
atically compared and recorded from culms, leaves and spike-like panicles. For lack of specimens, Phleum iranicum and
the two hybrid species are not included in the key. The characters and their states were so precisely defined that only 13
of them were needed to single out each taxon in the key. Detailed descriptions of all taxa based on the entire range of
20 characters are provided to aid identification.
Keywords: DELTA; identification; morphology; Maillea;Phalaris;Phleum;Pseudophleum
Introduction
The genus Phleum L. (Poaceae: Pooideae: Aveneae) com-
prises 16 species and two notho-species (Phleum ×
brueggeri K. Richt. and Phleum ×viniklarii Röhlena;
www.theplantlist.org; www.tropicos.org/names). The
chief centre of geographical distribution of the species is
the Mediterranean basin but some of them are distributed
widely in various habitats in northern Europe, Asia and
North and South America (Argentina and Chile) (Roshe-
vits 1980; Tzvelev 1983; Joachimiak and Kula 1997; Ste-
wart et al. 2011). Although none of the species is among
the globally important grain crops, a few perennial spe-
cies (e.g. Phleum pratense,Phleum alpinum,Phleum
phleoides) are grown as fodder and pasture plants in
numerous countries (Tzvelev 1983;Doğan 1991; Clayton
et al. 2002; Jefferson 2005; Smoliak et al. 2005; Stewart
et al. 2011). Phleum alpinum subsp. rhaeticum proved
useful for ecological restoration at high altitudes (Romani
et al. 2009), and P. pratense is commonly seeded for pas-
tures and hay and for ecological rehabilitation of dis-
turbed sites (Frank and Stephen 1983).
Outside its original Eurasian range, P. pratense has
now expanded to most suitable habitats around the
world. Accordingly, many agricultural studies have been
initiated to better understand its ecological and agricul-
tural impact in displacing native grasses (Stewart et al.
2005).
The identification of Phleum species and infra-speci-
fic taxa has been fraught with difficulties for a variety of
reasons: (i) the recognition of two or more infra-specific
taxa within several Phleum species and their perpetual
movement to and from different species of Phleum and
to other genera by various authors, (ii) the hierarchical
status of most infra-specific taxa was continually subject
to change and taxonomic controversy, (iii) the frequent
existence of several forms of the same species with dif-
ferent ploidy levels as in P. pratense, and (iv) the genus
has a checkered taxonomic history so that most of the
species were repeatedly moved to and from several other
genera (Achnodon Link, Chilochloa P. Beauv., Maillea
Parl., Phalaris L., Pseudophleum Doğan) by numerous
authors. For instance, Doğan (1982) established a new
monotypic genus [Pseudophleum gibbum (Boiss.)
Doğan] based on Phleum gibbum Boiss. This genus was
accepted by some subsequent authors (Doğan 1985;
Tzvelev 1989; Valdés and Scholz 2006,2009; Soreng
et al. 2015), and not others (e.g. Clayton and Renvoize
1986; Kellogg 2015), but has DNA support (Boudko
2014). Similarly, Phleum crypsoides (d’Urv.) Hack. was
originally placed in Phalaris and subsequently removed
to Maillea (Tzvelev 1989), before it was settled in its
currently accepted position in Phleum (Clayton and Ren-
voize 1986; Kellogg 2015; Soreng et al. 2015) and as a
separate section of Phleum by Horn af Rantzen (1946)
and Doğan (1985). A survey in the two sites of botanical
nomenclature (www.tropicos.org/NameSearch.aspx) and
(www.theplantlist.org/tpl/record/kew) showed that more
than 250 synonyms with no fewer than 17 generic names
are currently on record for the 16 Phleum species and
their sub-ordinate taxa.
The most comprehensive key to the species of
Phleum is that provided by Doğan (1991), although it
*Corresponding author. Email: elgazzar_adel@hotmail.com
© 2016 Dipartimento di Biologia, Università di Firenze
Webbia: Journal of Plant Taxonomy and Geography, 2016
Vol. 71, No. 1, 25–35, http://dx.doi.org/10.1080/00837792.2015.1119955
Published online 13 Jan 2016
does not include P. gibbum Boiss., the more recently
described Pseudophleum anatolicum Doğan, Behçet &
A. Sinan (Doğan et al. 2015) and the two notho-species.
Other keys include only the species found in local or
regional floras (e.g. Bor 1970; Humphries 1980; Tzvelev
1983;Doğan 1985; Barkworth 2007). These traditional
keys are manually constructed and exhibit some pitfalls
that lessen their practicality and reduce confidence in
their results. For example, in most of these keys the first
couplet is defined by the unobservable character “plants
annual versus perennial”, which renders further progress
in the key either impossible or shrouded with uncer-
tainty. In addition, the diagnostic character-states of con-
trasted entries in the same couplet are often not strictly
comparative or expressed by overlapping ranges of
value. Individual entries are frequently based on single
and/or vaguely defined characters. It is generally
acknowledged that some characters change considerably
among individual plants of the same taxon with variation
in habitat conditions or with the level of ploidy such as
“plants scapose versus lightly-densely caespitose”,“culm
erect versus geniculately ascending”, shoot height and
length of spikes (Callaghan 1974; Cenci et al. 1984; Cio-
sek et al. 2003). However, such unstable characters con-
tinued to feature prominently in most available keys.
This article is an attempt to depart from the choices
of ambiguous characters and character definitions and to
benefit from the inherent advantages of the well-tested
and highly efficient computer programs in the construc-
tion of a conventional key to Phleum species and their
subspecies based on the widest possible range of varia-
tion in their characters recorded from as many specimens
of every taxon as might cover the widest area of its
range of geographical distribution. Among the numerous
programs designed for key construction, the program
suite DELTA was successfully applied to the identifica-
tion of various groups of plants, animals and micro-or-
ganisms and is currently used worldwide (Dallwitz 1993
onwards; Dallwitz et al. 2000, 2010; Dallwitz and Paine
2005).
Material and methods
The present study is based on material obtained on loan
from the herbaria of the Botany Department, Faculty of
Science, Cairo University (CAI), the Royal Botanic Gar-
dens, Kew (K), the Missouri Botanic Gardens (MO),
Institute of Botany, University of Vienna (WU) and Nat-
ural History Museum of Florence (FI); acronyms are
according to Holmgren et al. (1990). Collection data of
all specimens are given in Appendix 1. The number of
specimens representing each taxon ranged from one (for
P. gibbum) to 25 (for P. alpinum subsp. alpinum)in
order to cover the widest possible range of its geographi-
cal distribution.
Identification of all specimens was scrutinized with
the aid of appropriate local and regional floras (Bor 1970;
Humphries 1980; Tzvelev 1983;Doğan 1985; Feinbrun-
Dothan 1986; Cope 2005; Barkworth 2007). Further con-
firmation of the plants’identity was carried out by match-
ing with detailed descriptions in the Grass Base
(www.kew.org/data/grasses-db).
For purposes of the present study, only the names
regarded as valid in (www.theplantlist.org/browse/A/)
and (www.tropicos.org/names) were accepted; except for
P. gibbum, synonyms and illegitimate and invalid names
are omitted. The accepted names of the species and their
subspecies together with full author citations are
arranged in alphabetical sequence as follows; taxa for
which no material was available are asterisked.
Phleum alpinum L., Sp. Pl. 1: 59. 1753
Two subspecies are recognized:
Phleum alpinum L. subsp. alpinum
Phleum alpinum L. subsp. rhaeticum Humphries, Bot.
J. Linn. Soc. 76: 339. 1978
Phleum arenarium L., Sp. Pl. 1: 60. 1753
Phleum bertolonii DC., Catal. Pl. Hort. Monsp. 132.
1813
(≡)Phleum pratense L. subsp. bertolonii (DC.) Bornm.,
Bot. Jahrb. 61, Beibl. 140: 157. 1928
Phleum boissieri Bornm., Magyar Bot. Lapok 11: 20.
1912
Phleum crypsoides (D’Urv.) Hackel, Bull. Soc. Bot.
Franc. 39: 274. 1892
Bas.: Phalaris crypsoides d’Urv., Mém. Soc. Linn. Paris
1: 263. 1822
Two subspecies are recognized:
Phleum crypsoides (D’Urv.) Hackel subsp. crypsoides
Phleum crypsoides (D’Urv.) Hackel subsp. sardoum
(Hackel ex Franch.) Horn af Rantzien, Bot. Not. 1946,
p. 370. 1946
Bas.: Phleum sardoum Hack. ex Franch., Bull. Soc. Bot.
France 39: 274. 1892
Phleum echinatum Host, Gram. Austr. 3: 8. 1805
Phleum exaratum Griseb., Spic. 2: 463. 1844
Two subspecies are recognized:
Phleum exaratum Griseb. subsp. exaratum
Phleum exaratum Griseb. subsp. aegaeum (Vierh.) W.
Greuter, Boissiera 13: 180. 1967
Bas.: Phleum arenarium L. subsp. aegaeum Vierh., Verh.
Zool-bot. Ges., Wien 69: 304. 1919
Phleum gibbum Boiss. Diagn. Pl. Orient, ser. 1, 5:
69–70. 1844
26 A. El-Gazzar et al.
Bas.: Pseudophleum gibbum (Boiss.) Dogan. Notes R.
Bot. Gard. Edinburgh, 40(1): 77. 1982
Phleum himalaicum Mez, Repert. Spec. Novi
Regni. Veg. 17: 293. 1921
Phleum hirsutum Honckeny, Verzeichn. Gewachse.
Deutschl. 1: 183. 1782
Phleum iranicum Bornm. & Gauba, Repert, Spec. Novi
Regni Veg. 47: 127. 1939*
Phleum montanum K. Koch, Linnaea 21: 383. 1848
Phleum paniculatum Hudson, Fl. Angl. (Hudson). 23.
1762
Phleum phleoides (L.) Karsten, Deutsche Fl. (Karsten).
374. 1881
Bas.: Phalaris phleoides L. Sp. Pl. 1: 55. 1753
Phleum pratense L., Sp. Pl. 1: 59. 1753
Two subspecies are recognized:
Phleum pratense L. subsp. pratense
Phleum pratense L. subsp.vulgare Čelak. Prodr. Fl.
Böhmen. 38. 1867
Phleum subulatum (Savi) Asch. & Graebn., Syn.
Mitteleur. Fl. 2(1): 154. 1899
Bas.: Phalaris subulata Savi, Fl. Pis. 1: 57. 1798
Two subspecies are recognized:
Phleum subulatum (Savi) Asch. & Graebn. subsp.
subulatum
Phleum subulatum (Savi) Asch. & Graebn. subsp.
ciliatum (Boiss.) Humphries, Bot. Journ. Linn. Soc. 76:
339. 1978
Bas.: Phleum tenue (Host) Schrad. var. ciliatum Boiss.,
Fl. Orient. 5: 480. 1884
Phleum ×brueggeri K. Richt. Pl. Europ. 1: 37. 1889.
[P. alpinum L. × P. michelii All. (= P. hirsutum
Honckeny)]*
P. ×viniklarii Röhlena Vestn. Král. Ceské Spolecn., Tr.
Mat.-Prir. 1933(8): 11. 1934. [Phleum echinatum Host ×
P. pratense L.?]*
The 20 characters selected for key construction are
listed in Table 1. The lengths of leaf blades were mea-
sured from the longest leaf on every available specimen
of each taxon and expressed as a range covering the
most frequent values so as to take intra-specific variation
into account. Shape of the spike-like panicle was
expressed as a range of the length/width ratios of at least
50 inflorescences of each taxon. A few mature spikelets
were resuscitated by boiling in water, dissected sepa-
rately on slides and cleared by warming in 1% KOH
solution. Lengths of the glumes and the hairs on their
keels were measured with the aid of a calibrated eye-
piece micrometer. Characters 15 and 18 in Table 1 were
verified by comparison with the scanning electron micro-
graphs published by Doğan (1982,1988) and Scholz
(1990,1999). No differences in the two-state characters
were observed between specimens of the same taxon. An
automated non-indented conventional key to the 20 taxa
was constructed using the program suite DELTA (Dall-
witz et al. 2000).
Results
Observations
The plants are annual or perennial, with single to loosely
tufted culms ranging in height from c. 4 cm in the two
subspecies of P. crypsoides to nearly 150 cm in some
specimens of P.pratense. The culms may be erect or
geniculately ascending, with a clearly swollen base in
some species (e.g. P. bertolonii and P. pratense). Leaves
are not aggregated basally and non-auriculate. Leaf
blades are flat, linear and 2.5–30 cm long and 1–15 mm
wide; these measurements are in close general agreement
with those scored by Casler (2001). The ligule is a
translucent membrane with a truncate or obtuse apex.
Each culm ends in a compact solitary terminal spike-like
panicle, which is usually 0.8–18 cm long and 0.2–1.2 cm
wide, and ranges in shape between almost globular to
narrowly cylindrical. The spikelet is invariably laterally
compressed, with a single flower and consists of two
glumes which are usually equal in size and shape (Fig-
ure 1B), but may be slightly sub-equal (Figure 1C, D).
Glumes vary considerably in shape, length : width ratio,
and in the distribution and length of hairs on their outer
surface (Figure 1). Each glume is keeled, three-veined
and has a green centre and two membranous margins
that vary in width from one species to another. The
backs of both glumes are commonly straight (Figure 1A,
C), but may be strongly curved inwards to give the spi-
kelet an ellipsoid shape (Figure 1E), or distally gibbous
as in P. gibbum. The apex of each glume may be trun-
cate (Figure 1A, C, D) or tapering (Figure 1B, E).
Glume apex ends either in a conspicuous awn (Fig-
ure 1A–D, F) or in a minute mucro (Figure 1E) so that
it appears awnless. The awn may be as long as the
glume body or longer (Figure 1A) or much shorter (Fig-
ure 1C, D, 6); the observed variation in awn length cor-
responds closely with that illustrated by Kováts (1976)
and Kula et al. (2006). Based on the wide differences in
their length, the hairs on the outer surface of glumes
were categorized into two clearly separate groups: (i)
long hairs ranging in length from 150 μm to about
1200 μm (Figure 1A–C), and (ii) short hairs not longer
Webbia: Journal of Plant Taxonomy and Geography 27
than 20 μm. Both categories share the pointed tip.
Although short hairs are observable on glumes of all
Phleum taxa, the much longer type is characteristic of
the glumes of only some taxa. The two glumes enclose a
single floret, which consists of the delicately membra-
nous and translucent lemma and palea (except in P. gib-
bum), two or three stamens with versatile two-lobed
anthers, two lodicules and a single-celled gynoecium.
The glumes are consistently much longer than the lem-
mas (Figure 1), except in P. gibbum. Lemmas are
broadly oblong or ovate, less firm than the glumes, trun-
cate or obtuse in shape, densely ciliate, sparingly hairy
or glabrous, and always awnless. The palea is slightly
shorter than the lemma and ciliate along the keel. The
clavate micro-hairs with rounded tips observed by Scholz
(1999) on the lemmas of P. subulatum have been found
also on the lemmas of P.boissieri and the two sub-
species of P.exaratum. Anthers are 0.3 to 2.3 mm long.
The ovary is glabrous and the styles are fused or free to
their bases and end in two feathery stigmas. The caryop-
sis is ellipsoid to ovoid, small and contains compound
starch grains (Humphries 1980; Watson and Dallwitz
1992; Barkworth 2007).
The Key
The following is an un-edited version of the conven-
tional key to the 20 taxa of Phleum; all characters were
given the same value of reliability (5.0):
Characters: 20 in data, 19 included, 13 in key.
Items: 20 in data, 20 included, 20 in key.
Parameters: Rbase = 1.40 Abase = 2.00 Reuse = 1.01
Varywt = .80
Characters included: 1–12 14–20
Character reliabilities: 1–20, 5.0
1. Leaf blade length less than 2 cm ....................... 2
Leaf blade length 4–10 cm .................................. 3
Leaf blade length 12 cm or more ..................... 11
2(1). Hairs on glume back 150–1200 µm
long ........................ P. crypsoides subsp. sardoum
Figures 1. Morphological variation in glumes of Phleum species. (A) Phleum alpinum ssp. alpinum, truncate apex, long hairs on
midrib, awn nearly as long as glume.(B) Phleum hirsutum, tapering apex, long hairs on midrib and awn. (C) Phleum arenarium,
truncate apex, long hairs on midrib, awn much shorter than glume. (D) Phleum paniculatum, truncate apex, micro-hairs, awn much
shorter than glume. (E) Phleum subulatum ssp. subulatum, tapering apex, micro-hairs, awnless. (F) Phleum phleoides, tapering apex,
micro-hairs, awn much shorter than glume.
28 A. El-Gazzar et al.
Hairs on glume back up to 20 µm
long .................... P. crypsoides subsp. crypsoides
3(1). Awn as long as glume or longer...... P. echinatum
Awn shorter than glume......................................4
Awn absent ........................................................ 10
4(3). Leaf sheath inflated ............................................. 5
Leaf sheath not inflated....................................... 8
5(4). Glume apex truncate ...........................................6
Glume apex tapering ........................... ................7
6(5). Hairs on glume back 150–1200 µm long; culm
base swollen ..................................... P. arenarium
Hairs on glume back up to 20 µm long; culm
base not swollen ............................ P. paniculatum
7(5). Spikelet adnate to rachis; hairs on glume
subapical margin absent; culm basenot
swollen ............................................ P. himalaicum
Spikelet free from the; hairs on glume subapical
margin present; culm base rachis........P. hirsutum
8(4). Hairs on glume subapical margin pre-
sent ......................... P. exaratum subsp. exaratum
Hairs on glume subapical margin absent............9
9(8). Clavate hairs on lemma present
..................................P. exaratum subsp. aegaeum
Clavate hairs on lemma absent ........ P. montanum
10(3). Leaf sheath inflated; spike-like panicle
length/width ratio 15 or more; spikelet free from
the rachis; rachis extension present
............................. P. subulatum subsp. subulatum
Leaf sheath not inflated; spike-like panicle
length/width ratio 4–11; spikeletadnate to rachis;
rachis extension absent.......................... P. gibbum
11(1). Awn as long as glume or longer ......................12
Awn shorter than glume....................................13
Awn absent ..............P. subulatum subsp. ciliatum
12(11). Awn ciliate .............. P. alpinum subsp. rhaeticum
Awn not ciliate ........... P. alpinum subsp. alpinum
13(11). Leaf sheath inflated...........................................14
Leaf sheath not inflated ................................... 15
14(13). Spikelet adnate to rachis; glume apex truncate;
hairs on glume subapicalmargin present; ligule
truncate.....................P. pratense subsp. pratense
Spikelet free from the rachis; glume apex taper-
ing; hairs on glume subapicalmargin absent;
ligule acute-obtuse ............................. P. boissieri
15(13). Ligule acute-obtuse; rachilla extension absent16
Ligule truncate; rachilla extension pre-
sent............................ P. pratense subsp. vulgare
16(15). Spikelet adnate to rachis; culm base
swollen............................................. P. bertolonii
Spikelet free from the rachis; culm base not
swollen..............................................P. phleoides
Descriptions
The following is an un-edited version of the detailed
descriptions obtained from the DELTA run for the 20
Phleum taxa in terms of the 20 characters and their charac-
ter-states listed in Table 1 ; all taxa are in alphabetical order.
Phleum alpinum subsp.alpinum
Culm at least 8 cm long. Culm base not swollen. Spike-
like panicle length : width ratio 4–11. Leaf blade length
12 cm or more. Leaf sheath inflated. Ligule acute-obtuse.
Spikelet adnate to rachis. Rachilla extension absent.
Glumes equal. Glumes longer than lemma and palea.
Glume back straight. Glume apex truncate. Glume length
5.6 mm. Hairs on glume back 150–1200 μm long. Hairs
on glume subapical margin absent. Awn as long as
glume body or longer. Awn not ciliate. Globose projec-
tions on lemma surface absent. Clavate hairs on lemma
absent. Three stamens.
Table 1. List of 20 characters and their character-states as recorded comparatively for 20 taxa of Phleum and used in key construc-
tion.
1. Culm/ 1. at least 8 cm long/ 2. up to 4 cm long/
2. Culm base/ 1. swollen/ 2. not swollen/
3. Spike-like panicle length/width ratio/ 1. 1.5–3/ 2. 4–11/ 3. 15 or more/
4. Leaf blade length/ 1. less than 2 cm/ 2. 4–10 cm/ 3. 12 cm or more/
5. Leaf sheath/ 1. inflated/ 2. not inflated/
6. Ligule/ 1. acute-obtuse/ 2. truncate/
7. Spikelet/ 1. adnate to rachis/ 2. Free from the rachis/
8. Rachilla extension/ 1. present/ 2. absent/
9. Glumes/ 1. equal/ 2. unequal/
10. Glumes/ 1. longer than lemma and palea/ 2. shorter than lemma and palea/
11. Glume back/ 1. straight/ 2. curved inwards/ 3. distally gibbous/
12. Glume apex/ 1. truncate/ 2. tapering/
13. Glume length/ mm/
14. Hairs on glume back/ 1. 150–1200 μm long/2. up to 20 μm long/
15. Hairs on glume subapical margin/ 1. present/ 2. absent/
16. Awn/ 1. as long as glume body or longer/ 2. shorter than glume body/ 3. absent/
17. Awn/ 1. ciliate/ 2. not ciliate/
18. Globose projections on lemma surface/ 1. present/ 2. absent/
19. Clavate hairs on lemma/ 1. present/ 2. absent/
20. Stamens/ 1. 3/ 2. 2/
Webbia: Journal of Plant Taxonomy and Geography 29
Phleum alpinum subsp.rhaeticum
Culm at least 8 cm long. Culm base not swollen. Spike-
like panicle length : width ratio 4–11. Leaf blade length
12 cm or more. Leaf sheath inflated. Ligule acute-obtuse.
Spikelet adnate to rachis. Rachilla extension absent.
Glumes equal. Glumes longer than lemma and palea.
Glume back straight. Glume apex truncate. Glume length
5.2 mm. Hairs on glume back 150–1200 μm long. Hairs
on glume subapical margin absent. Awn as long as
glume body or longer. Awn ciliate. Globose projections
on lemma surface absent. Clavate hairs on lemma absent.
Three stamens.
Phleum arenarium
Culm at least 8 cm long. Culm base swollen. Spike-like
panicle length/width ratio 4–11. Leaf blade length 4–
10 cm. Leaf sheath inflated. Ligule acute-obtuse. Spikelet
free from the rachis. Rachilla extension absent. Glumes
equal. Glumes longer than lemma and palea. Glume back
straight. Glume apex truncate. Glume length 2.8 mm.
Hairs on glume back 150–1200 μm long. Hairs on glume
subapical margin absent. Awn shorter than glume body.
Awn not ciliate. Globose projections on lemma surface
absent. Clavate hairs on lemma absent. Three stamens.
Phleum bertolonii
Culm at least 8 cm long. Culm base swollen. Spike-like
panicle length/width ratio 4–11. Leaf blade length 12 cm
or more. Leaf sheath not inflated. Ligule acute-obtuse. Spi-
kelet adnate to rachis. Rachilla extension absent. Glumes
equal. Glumes longer than lemma and palea. Glume back
straight. Glume apex truncate. Glume length 3.7 mm.
Hairs on glume back 150–1200 μm long. Hairs on glume
subapical margin absent. Awn shorter than glume body.
Awn not ciliate. Globose projections on lemma surface
absent. Clavate hairs on lemma absent. Stamens 3.
Phleum boissieri
Culm at least 8 cm long. Culm base not swollen. Spike-
like panicle length : width ratio 4–11. Leaf blade length
12 cm or more. Leaf sheath inflated. Ligule acute-obtuse.
Spikelet free from the rachis. Rachilla extension absent.
Glumes equal. Glumes longer than lemma and palea.
Glume back straight. Glume apex tapering. Glume length
3.6 mm. Hairs on glume back 150–1200 μm long. Hairs
on glume subapical margin absent. Awn shorter than
glume body. Awn not ciliate. Globose projections on
lemma surface absent. Clavate hairs on lemma present.
Three stamens.
Phleum crypsoides subsp. crypsoides
Culm up to 4 cm long. Culm base not swollen. Spike-
like panicle length : width ratio 1.5–3. Leaf blade length
less than 2 cm. Leaf sheath inflated. Ligule truncate. Spi-
kelet free from the rachis. Rachilla extension present.
Glumes equal. Glumes longer than lemma and palea.
Glume back straight. Glume apex tapering. Glume length
3.7 mm. Hairs on glume back up to 20 μm long. Hairs
on glume subapical margin absent. Awn shorter than
glume body. Awn not ciliate. Globose projections on
lemma surface absent. Clavate hairs on lemma absent.
Two stamens.
Phleum crypsoides subsp.sardoum
Culm up to 4 cm long. Culm base not swollen. Spike-
like panicle length : width ratio 1.5–3. Leaf blade length
less than 2 cm. Leaf sheath inflated. Ligule truncate. Spi-
kelet free from the rachis. Rachilla extension present.
Glumes equal. Glumes longer than lemma and palea.
Glume back straight. Glume apex tapering. Glume length
3.3 mm. Hairs on glume back 150–1200 μm long. Hairs
on glume subapical margin absent. Awn shorter than
glume body. Awn not ciliate. Globose projections on
lemma surface absent. Clavate hairs on lemma absent.
Two stamens.
Phleum echinatum
Culm at least 8 cm long. Culm base not swollen. Spike-like
panicle length : width ratio 1.5–3. Leaf blade length
4–10 cm. Leaf sheath inflated. Ligule acute-obtuse. Spike-
let adnate to rachis. Rachilla extension absent. Glumes
equal. Glumes longer than lemma and palea. Glume back
straight. Glume apex truncate. Glume length 3.5 mm. Hairs
on glume back 150–1200 μm long. Hairs on glume subapi-
cal margin present. Awn as long as glume body or longer.
Awn not ciliate. Globose projections on lemma surface
absent. Clavate hairs on lemma absent. Three stamens.
Phleum exaratum subsp.aegaeum
Culm at least 8 cm long. Culm base swollen. Spike-like
panicle length : width ratio 4–11. Leaf blade length 4–
10 cm. Leaf sheath not inflated. Ligule acute-obtuse. Spi-
kelet free from the rachis. Rachilla extension absent.
Glumes equal. Glumes longer than lemma and palea.
Glume back straight. Glume apex tapering. Glume length
3.2 mm. Hairs on glume back 150–1200 μm long. Hairs
on glume subapical margin absent. Awn shorter than
glume body. Awn not ciliate. Globose projections on
lemma surface absent. Clavate hairs on lemma present.
Three stamens.
Phleum exaratum subsp. exaratum
Culm at least 8 cm long. Culm base swollen. Spike-like
panicle length : width ratio 4–11. Leaf blade length 4–
10 cm. Leaf sheath not inflated. Ligule acute-obtuse. Spi-
kelet free from the rachis. Rachilla extension absent.
Glumes equal. Glumes longer than lemma and palea.
Glume back straight. Glume apex tapering. Glume length
30 A. El-Gazzar et al.
3.9 mm. Hairs on glume back 150–1200 μm long. Hairs
on glume subapical margin present. Awn shorter than
glume body. Awn not ciliate. Globose projections on
lemma surface absent. Clavate hairs on lemma present.
Three stamens.
Phleum gibbum
Culm at least 8 cm long. Culm base not swollen.
Spike-like panicle length : width ratio 4–11. Leaf blade
length 4–10 cm. Leaf sheath not inflated. Ligule acute-
obtuse. Spikelet adnate to rachis. Rachilla extension
absent. Glumes unequal. Glumes shorter than lemma and
palea. Glume back distally gibbous. Glume apex taper-
ing. Glume length 2.9 mm. Hairs on glume back up to
20 μm long. Hairs on glume subapical margin absent.
Awn absent. Awn not ciliate. Globose projections on
lemma surface present. Clavate hairs on lemma absent.
Three stamens.
Phleum himalaicum
Culm at least 8 cm long. Culm base not swollen.
Spike-like panicle length : width ratio 4–11. Leaf blade
length 4–10 cm. Leaf sheath inflated. Ligule acute-ob-
tuse. Spikelet adnate to rachis. Rachilla extension
absent. Glumes equal. Glumes longer than lemma and
palea. Glume back straight. Glume apex tapering.
Glume length 3.5 mm. Hairs on glume back 150–
1200 μm long. Hairs on glume subapical margin
absent. Awn shorter than glume body. Awn not ciliate.
Globose projections on lemma surface absent. Clavate
hairs on lemma absent. Three stamens.
Phleum hirsutum
Culm at least 8 cm long. Culm base swollen. Spike-like
panicle length : width ratio 4–11. Leaf blade length 4–
10 cm. Leaf sheath inflated. Ligule acute-obtuse. Spikelet
free from the rachis. Rachilla extension absent. Glumes
equal. Glumes longer than lemma and palea. Glume back
straight. Glume apex tapering. Glume length 4.8 mm.
Hairs on glume back 150–1200 μm long. Hairs on glume
subapical margin present. Awn shorter than glume body.
Awn not ciliate. Globose projections on lemma surface
absent. Clavate hairs on lemma absent. Three stamens.
Phleum montanum
Culm at least 8 cm long. Culm base swollen. Spike-like
panicle length : width ratio 4–11. Leaf blade length 4–
10 cm. Leaf sheath not inflated. Ligule acute-obtuse. Spi-
kelet free from the rachis. Rachilla extension absent.
Glumes equal. Glumes longer than lemma and palea.
Glume back straight. Glume apex tapering. Glume length
3.2 mm. Hairs on glume back 150–1200 μm long. Hairs
on glume subapical margin absent. Awn shorter than
glume body. Awn not ciliate. Globose projections on
lemma surface absent. Clavate hairs on lemma absent.
Three stamens.
Phleum paniculatum
Culm at least 8 cm long. Culm base not swollen. Spike-
like panicle length : width ratio 4–11. Leaf blade length
4–10 cm. Leaf sheath inflated. Ligule acute-obtuse. Spi-
kelet free from the rachis. Rachilla extension absent.
Glumes equal. Glumes longer than lemma and palea.
Glume back straight. Glume apex truncate. Glume length
1.9 mm. Hairs on glume back up to 20 μm long. Hairs
on glume subapical margin absent. Awn shorter than
glume body. Awn not ciliate. Globose projections on
lemma surface absent. Clavate hairs on lemma absent.
Three stamens.
Phleum phleoides
Culm at least 8 cm long. Culm base not swollen. Spike-
like panicle length : width ratio 4–11. Leaf blade length
12 cm or more. Leaf sheath not inflated. Ligule acute-ob-
tuse. Spikelet free from the rachis. Rachilla extension
absent. Glumes equal. Glumes longer than lemma and
palea. Glume back straight. Glume apex truncate. Glume
length 2.9 mm. Hairs on glume back 150–1200 μm long.
Hairs on glume subapical margin absent. Awn shorter
than glume body. Awn not ciliate. Globose projections
on lemma surface absent. Clavate hairs on lemma absent.
Three stamens.
Phleum pratense subsp.pratense
Culm at least 8 cm long. Culm base swollen. Spike-like
panicle length : width ratio 4–11. Leaf blade length
12 cm or more. Leaf sheath inflated. Ligule truncate.
Spikelet adnate to rachis. Rachilla extension absent.
Glumes equal. Glumes longer than lemma and palea.
Glume back straight. Glume apex truncate. Glume length
6.2 mm. Hairs on glume back 150–1200 μm long. Hairs
on glume subapical margin present. Awn shorter than
glume body. Awn not ciliate. Clavate hairs on lemma
absent. Three stamens.
Phleum pratense subsp. vulgare
Culm at least 8 cm long. Culm base swollen. Spike-like
panicle length : width ratio 4–11. Leaf blade length
12 cm or more. Leaf sheath not inflated. Ligule truncate.
Spikelet adnate to rachis. Rachilla extension present.
Glumes equal. Glumes longer than lemma and palea.
Glume back straight. Glume apex truncate. Glume length
4.9 mm. Hairs on glume back 150–1200 μm long. Hairs
on glume subapical margin absent. Awn shorter than
glume body. Awn not ciliate. Globose projections on
Webbia: Journal of Plant Taxonomy and Geography 31
lemma surface absent. Clavate hairs on lemma absent.
Three stamens.
Phleum subulatum subsp. ciliatum
Culm at least 8 cm long. Culm base not swollen. Spike-
like panicle length : width ratio 15 or more. Leaf blade
length 12 cm or more. Leaf sheath inflated. Ligule acute-
obtuse. Spikelet free from the rachis. Rachilla extension
present. Glumes equal. Glumes longer than lemma and
palea. Glume back curved inwards. Glume apex tapering.
Glume length 2.7 mm. Hairs on glume back 150–
1200 μm long. Hairs on glume subapical margin absent.
Awn absent. Awn not ciliate. Globose projections on
lemma surface present. Clavate hairs on lemma present.
Three stamens.
Phleum subulatum subsp.subulatum
Culm at least 8 cm long. Culm base not swollen. Spike-
like panicle length/width ratio 15 or more. Leaf blade
length 4–10 cm. Leaf sheath inflated. Ligule acute-ob-
tuse. Spikelet free from the rachis. Rachilla extension
present. Glumes equal. Glumes longer than lemma and
palea. Glume back curved inwards. Glume apex tapering.
Glume length 4.3 mm. Hairs on glume back up to 20 μm
long. Hairs on glume subapical margin absent. Awn
absent. Awn not ciliate. Globose projections on lemma
surface present. Clavate hairs on lemma absent. Three
stamens.
Discussion
Glume morphology proved to be an essential tool in the
circumscription and identification of specific and infra-
specific taxa within Phleum. It provides most of the
characteristics that have a diagnostic value in the recog-
nition of all species and subspecies.
The recent phylogenetic findings by Boudko (2014)
and Soreng et al. (2015) support the isolation of P. gib-
bum in the genus Pseudophleum by Doğan (1982). How-
ever, P. gibbum was included in the present key because
the morphological differences between this species and
the rest of Phleum are not greater than those between
any other species and the rest. In fact, the differences
between the two subspecies of P. subulatum and other
Phleum species are much greater than those between
P. gibbum and the rest, so that P. subulatum ought to
have been the likely candidate for separation from
Phleum. Nevertheless, P. subulatum was not removed
from Phleum because recent studies by Eisa (2012) and
El-Gazzar et al. (2013) have indicated a close relation-
ship between it and the other two species in Phleum-sec-
tion Achnodon (P. exaratum and P. boissieri).
The computer-generated key provided in the present
study avoids the numerous shortcomings of previous
manually constructed keys. It is strictly comparative in
that alternative entries of the same couplet are diagnosed
by the contrasted character-states. The states of all char-
acters are clearly defined with the maximum possible dif-
ferences between them so that the user can easily decide
which character-state applies to the unknown specimens.
Among the numerous advantages inherent in the program
suite DELTA is the production of detailed descriptions of
individual taxa in terms of the entire set of recorded
characters. The list of parameters preceding the key indi-
cates that of the 20 characters recorded for the taxa only
13 were sufficient to single out every one of the 20 taxa.
A surplus of seven characters in these comparative
descriptions serves the important function of confirming
the results of using the key.
Only accepted names appear in the key and descrip-
tions. Full lists of synonyms may be obtained from the
two websites [www.tropicos.org/NameSearch.aspx;
www.theplantlist.org/tpl/record/kew]. Phleum iranicum
Bornm. & Gauba (an annual endemic to Iran) and the
two notho-species (P. ×brueggeri K. Richt. and
P. ×viniklarii Röhlena) are missing in the present key
because of a lack of specimens. Whether P. gibbum and
its close relative Pseudophleum anatolicum (Doğan et al.
2015) are kept within Phleum or treated as a separate
genus, they can be added (together with P. iranicum and
the two notho-species), to the present data set and
included in an expanded version of the present key when-
ever sufficient material becomes available. The data
matrix used in the present study might also be expanded
to accommodate any additional characters that might be
needed to distinguish between all the taxa in the new key.
Acknowledgements
Thanks are due to the Keepers of the herbaria of the Royal
Botanic Gardens, Kew, the Missouri Botanic Gardens, Institute
of Botany, University of Vienna and Natural History Museum
of Florence for the generous loans of specimens.
Disclosure statement
No potential conflict of interest was reported by the authors.
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Appendix 1
Collection data of specimens of Phleum species and subspecies
in the following order: taxon. Country, locality; collector, col-
lector’s number if available; (herbarium acronym); accession
number if available; date of collection.
Phleum alpinum L.subsp alpinum
Austria, Tirol, Obergurgel; Nabil El Hadidi (CAI);
26.7.1958; Austria, Jämtland, Åre, Saasarna, Getryggen; E.
Folkesen; (CAI); 21.7.1960; Austria, Jämtland, Åre, Benfjället;
Webbia: Journal of Plant Taxonomy and Geography 33
E. Folkesen; (CAI); 3.8.1960; Austria, Jämtland, Åre, Tage-
fors; E. Folkesen; (CAI); 6.7.1968; Switzerland, La Linnaea,
Burg St. Pierre; A.M. Migahid & A.H. Montasir; (CAI); June-
August 1936; Hungary; S. Kupčok; (CAI); 4.7.1994; Sweden,
Bjiorkliden; Robert Lamm; (CAI); 22.6.1921; Sweden, Abisko;
M. Ayyad; (CAI); July 1958; Italy, Vallistellina, prope Bormio;
C. Camperio; (CAI); 16 August 1904; Canada, W.G.Dore,
(MO), 1826348, 1948; Guatemala, M. D. Nyj, (MO),
04819005, 1998; Romania, E. I. Nyarady, (MO), 1929; Fin-
land, O. Kyyhkynen, (MO), 1087985, July 1918 Finland, Hel-
sinki, Bjorn Pederely, (MO), 2785807, August 1973. Argentina,
(MO), 1164434, March 1936. Brazil, Harry Smith, (MO),
1641704, August 1950. Norway, J.E. Elsley, (MO), August
1966. Alaska, S. Studebaker and C.L. Parker, (MO), 5970530,
July 2006. Argentina, F. Roig et al., (MO), 3504060, January
1978. Colorado, J.R Churchill, (MO), 1057104, July 1912.
Wyoming, Louis Williams, (MO), 1046733, November 1933.
Mexico, C.A. Puiplus, (MO), 3047874, September 1907. Ice-
land, A.E. Skjot Pederson, (MO), 3473337, June 1924. Green-
land, J. Eugenius, (MO), 1091322, July 1934. Italy, C.
Camperio, (MO), August 1904. Arizona, C.F. Cox, (MO),
1120967.
Phleum alpinum L. subsp. rhaeticum Humphries
Austria, Oberösterreich, Dachstein, Gipfelbereich, G. Klee-
sadl 485, (CAI), 18.8.1995. Austria, Vienna, Manfred A. Fisher
(WU), 5075, 1996. Austria, Joul Breidler, (WU), 1868. Afgha-
nistan, Lerpow, (WU), 1935. Greece, Daneil Rape, (WU),
4362, 1934. Iran, Abdallah Riaze, (WU), 2346, 1946. Austria,
Joul Breidler, (WU), 6342, 1871. Greece, Manfred A. Fisher,
(WU), 991, 1987. Austria, Vienna, Manfred A. Fisher, (WU),
5077, 1996. Afghanistan, Talal Rouh, (WU), 2513, 1947. Italy,
L. Boulos, (WU), 37645, 1988. Austria, Lauren, (WU), 845,
1979.
Phleum arenarium L.
Germany, Insula Hiddensae, B. Lorenz, (CAI), July 1913.
England, Burnham, Somerset, S.H. Thompson, (CAI), 1889.
England, Ogaden, Glomorgan, S.H. Thompson, (CAI), 29
August 1920. England, Burnham, N. Somerset, H.S. Thomp-
son, (CAI), 7 June 1923. England, Mildenhall, Suffolk, J.S.
Henslow, (CAI), 3 June 1829. England, Liverpool, W. Wilson,
(CAI), June 1822. England, A.E. Lomax, (MO), 3047917, May
1885. Sweden, Alvaret, Vickleby, Öland, Tovart Laurent,
(CAI), 10.8.1943. Denmark, P. Pedersen & B. Öllgaard, (CAI),
26.6.1969. Himalaya, R.R. Stewart, (MO), 1262473, April
1938. Pakistan, B. Dickore, (MO), 5660882, 1995. Pakistan, A.
Mombucher, (MO), 2885102, June 1890. Gotland, Erik
Asplund, (MO), 937861, June 1926. Netherlands, L.Y. Westra,
(MO), 4048873, 1959. Denmark, L. Br. Holm-Nielsen, (MO),
2196522, 1967. Morocco, (MO), 2307005, 1974. Netherland,
Friedrich Zimmerman, (WU), 2427, June 1906. USA, Dr. Grie-
wank, (WU), 2498, July 1982. Palestine, J. Bronmuller, (WU),
1606, June 1897. Greece, Chr. Leonis, (WU), 91, May 1898.
Germany, K.H. Rechinger, (MO), 1743664, 1958. Italy, Adr.
Fiori, (WU), 2718, May 1913. England, A.E.Lomax, (MO),
3047917, June, 1885.
Phleum bertolonii (DC) Bornm.
France, dép. Gard, Les Angles, J.C Ledoux, (CAI),
16.7.1974. France, P. Martin, (MO), 39225771, June 1981.
Denmark, Benjamin Öllgaard, (MO), 2765157, 1962. USA, B.
Richard, (WU), 27615, 1956. Denmark, (K), 6539820, 1967.
Italy, Brane, (Fl), 714362, 1987. Germany, Gabriel Stroby,
(WU), 1990. Germany, Jichler, (MO), 1833420, 1945. France,
Lauren De, (Fl), 234165, 1987. Italy, (K), 4381003, 1976.
USA, Manfred A. Fischer, (Fl), 32415, 1998.
Phleum boissieri Hochst. ex Griseb.
Syria, J. Bornmuller, (WU), 1910. Iran, Pahat, (WU),
10165, 1959. Turkey, Henderson and Stainton, (K), 5570,
1960. Turkey, Davis and Hedge, (K), 28737, 1957. Greece, C.
H. Doris, (K), 4308, 1942. Iraq, J.B. Gillett, (K), 11584, 1948.
Iran, Walter Koeltz, (K), 1940. Iran, Laranhabs, (K), 10293,
1968.
Phleum crypsoides (D’Urv.) Hackel subsp. crypsoides
France, Weilber, (WU),10742, 1931. Greece, (K), 2305,
1962. Italy, Baldoran, (K), 1953. Greece, Heldreich, (Fl),
66457, 1964. Greece, J. Prudhomme, (WU), 15002, 1968.
Phleum crypsoides (D’Urv.) Hackel subsp. sardoum (Hackel)
Horn af Rantzien
Sardaigne, (K), 149, 1881. Sardaigne, A. E. Willmott, (K),
149, 1881. Sardaigne, R. D. Meikle, (K), 2305, 1962. Sar-
daigne, S.C. Ashley, (K), 1074, 1931.
Phleum echinatum Host
Kroatia, Fr. Petter, (MO), 1025660, 1868. Germany, Jichler,
(MO), 3047970, 1868. Germany, Jichler, (MO), 1870. Italy,
(MO), 3047975, 1897. Koroatia, L. Grossu and A. Kneueker,
(MO), 2882819, 1902. Italy, E. Preissauz, (MO), 1025659,
1911. Germany, Gabriel Stroby, (WU), June 1874. Italy, Erwin
Janchen, (WU), 2278, 1906. Italy, W. Noeiter, (MO), 3047977,
1844. Italy, Antonio Baldacci, (MO), 3047974, 1895.
Phleum exaratum Hoschst. in Boiss. subsp. aegaeum (Vierh.)
W. Greuter Crete Island, Moland and Tarek, (MO), 4376580,
1964. Crete Island, (MO), 4376580, 1964. Crete Island, Rafaeil
Donas, (K), 4375946, 1978. Crete Island, J. Prudhomme, (K),
4539087, 1962. Crete Island, (K), 3419836, 1964. Crete Island,
(K), 3652870, 1988.
Phleum exaratum Hoschst. in Boiss., subsp. Exaratum
Syria, Gunnar Samuelsson, (K), 0171573, 1933. Iraq, M.E.
Boose, (K), 537, 1958. Turkey, Davis and Dodds, (K), 01715,
1952. Serbia. L.S. Winjrcie, (WU), 1896. Greece, Heldreich,
(WU). Greece, J. Bornmüller, (WU), 1909. Greece, L. Adamo-
vic, (WU), 1943. Syria, K.H. Rechinger, (MO), 4376580, June
1964. Turkey, Hardel, (WU), 450, 1889. Syria, Gunnar
Samuelsson, (WU), 4959, 1933. Lebanon, Breidly, (WU),
2006. Afghanistan, (K), 285, 1969. Iran, M. Jacobs, (WU),
6769, 1963. Iraq, Raui & Namik, (K), 1959.
Phleum gibbum Boiss.
Turkey, B. Balansa, (CAI), July 1857.
Phleum himalaicum Mez
India, Himalaya, V. Jacquemont 297, (K), 10.5.1848. India,
Himalayia, D. Khan, (Fl), 2341, 1954. India, Himalayia, D.
Khan, (Fl), 2343, 1954. Pakistan, A. Hamid, (WU), October
1972. Iran, Dughes, (Fl), 1962. Pakistan, L. Christopher, (MO),
4308276, May 1987. Afghanistan, M. Robert, (WU), 1432,
1982.
Phleum hirsutum Honckeny
Germany, Algäu, K. Richter, (CAI), no date. Germany, K.
Richter, (CAI), 21.7.1886. Switzerland, R. Barbazat, (CAI),
15.8.1958. Cyperus, Carpasia, A. Margrissin, (CAI), July 1925.
Montenegro, W. Guterman, (WU), 5056, 2007. France, Quadr-
ual, (WU), 4785. France, J. Prudhomme, (WU), 4955, 1960.
Poland, B. Pawowski, (MO), 1120400, August, 1932. Italy, E.
A. Huetdy, (MO), 3847999, 1855. Austria, Manfred A. Fischer,
(WU), 5075, 1996. Austria, (MO), 908550, June 1888. Roma-
nia, Prodan, (MO), 1101882. Romania, A. Avn Degen, (MO),
2709633, 1905.
Phleum montanum C. Koch
Lebanon, Vivi Täckholm, (CAI), June 1954. Russia, E.N.
Gym, (CAI), 22.7.1930. Georgia, S. Renvoize, (MO), 5864153,
July 2004. Italy, Jalis, (MO), 705552, 1872. Spain, A. Gabel-
lew, (MO), 1639164, 1932. Syria, Gunnar Samuelsson, (MO),
2363483, May 1939. Turkey, E.L. Nielsen, (MO), 1837893,
34 A. El-Gazzar et al.
September 1962. Romania, Peter fi, (MO), 915444, May 1918.
Georgia, Otar Abd Aladze, (MO), 5936678, July 2004.
Phleum paniculatum Huds.
Austria, umgebung von Basl, Reichenbach f., (CAI), no
date. Germany, Hessen, Nassau, A.W. Peipers, (CAI), July
1907. Germany, C. Baenitz and Nordu, (MO), 909767, 1865.
Germany, (MO), 5988888, 1890. Germany, F. Persinger, (MO),
1857. Russia, R.F. Hohenachen, (MO), 3048090, 1843. Russia,
A. Gordjogin, (MO), 1660003, 1916. Russia, V. Goloskokov,
(MO), 5683392, 1963. China, K. Yao, (MO), 3629803, May
1984. China, Dai Tainlan, (MO), 4735142, May 1958. China,
A.N. Steward, (MO), 904192, May 1922. China, Taiyang Ping
and Q. S. Wang, (MO), 4511940, September 1981. Georgia,
Gagnize, (MO), 04929862, 1997.
Phleum phleoides (L.) Karsten
Sweden, Furulund, Skåne, A. El-Gazzar, (CAI), August
1976. Denmark, Peder Pedersen & Benjamin Öllgaard, (CAI),
15.7.1960. Germany, Köln, Bernh. Lorenz, (CAI), January
1895. Germany, Schuster, (CAI), July 1910. Switzerland,
Gland, (CAI), October 1935. Russia, A. Muselbeou 1606,
(CAI), June 1910. Austria, La Linnaea, Burg St. Pierre, A.M.
Migahid & A.H. Montasir, (CAI), June-August 1936. Slovenia,
J. Podpera, (MO), 930939, May 1926. Iran, N.L. Bor, (MO),
5649549, February 1948. Russia, (MO), 5189759, July 1994.
China, Da Fang, (WU), 1930. France, A. Kneueker, (MO),
2885103, June 1899. Finland, K. E.. Hirn, (MO), 864766,
1894. Poland, R. Kobendza, (MO), 1120401. April 1932. Geor-
gia, Nikoloz Lachashvili, (MO), 5965132, May 2005. Norway,
Hartvig Johnsenn, (WU), 1910. France, (MO), May 1969. Mor-
occo, Tizi-n-Treiten, (MO), 2306482, June 1974.
Phleum pratense L. subsp. pratense
Sweden, Öland, Gårdby, Ove Almborn, (CAI), 10.7.1935.
Denmark, L.B. Holm, (MO), 2191029, July 1966. Germany,
Franz Petji, (MO), 380, August 1900. Canada, J. A. Calder &
R. L. Taylor, (MO), 1995631, July 1964. New Mexico, Brian
Reif, (MO), 04856482, August 2002. England, Wurzell, (MO),
2345494, July 1973. Australia, L. A. Johnson, (MO), 1622631,
January 1951. USA, Colorado, Robert Merril & Robert H. Gar-
vey, (MO), 04840116, October 2004. China, H. R. Shan Lai &
Shan, (MO), 04510040, September 1996. Iceland, A. E. Skjot
Pedersen, (MO), 3473338, July 1924. Chile, M. R. Espinosa,
(MO), 1211759, February 1940. Japan, Koji Yonekura et al.,
(MO), 6050271, June 1993. Finland, A. Oswald Kihlman,
(MO), 864772, August 1864. Canada, J. Bousseau, (MO),
982042, July 1962. Canada, R. C. Hosie & E.O. Hughes,
(MO), 17103662, July 1938. Argentina, D. Swallen and R. De
Barba, (MO), 157439, January 1947. Czechoslovakia, Jan
Bohus Slavek, (MO), 3907345, July 1991. Georgia, M. Khut-
sishvili, (MO), 5941668, September 2006. Japan, Voichi
Tateishi & Takahide Kurosasa, (MO), 4254505, August 1991.
Phleum pratense L. subsp. vulgare
Sweden, Jämtland, Frösön, Vagled, E. Folkeson, (CAI),
12.8.1952. Sweden, Bjorkliden, Robert Lamm, (CAI),
22.6.1921. Sweden, Vedervåg, (CAI), 17.6.1929. Sweden,
Skåne, A. Torlöw, 15.7.1936. Sweden, Erik Evers, (CAI),
10.7.1960. Sweden, Öland, Södra Alvaret, Alby, E. Folkeson,
1.7.1972. Sweden, H. Hylander, (CAI), 29.6.1935. Denmark, P.
Holm, Ivan Nielsen & P. Pedersen, (CAI), 22.8.1966. Austria,
La Linnaea, Burg St Pierre, A.M. Migahid & A.H. Montasir,
(CAI), June–August 1936. Germany-West, Oberbayern, Erich
Albertshofer, (CAI), 7.7.1972. Italy, Piemonte, A. Johnson &
E. Rowsetti, (CAI), 24.6.1921. Spain, N.P. Blat & J. Quessa
5340, (CAI), 30.5.1966. Hungary, Samuel Kupčok, (CAI),
18.6.1908. Russia, A.E. Androponov, (CAI), 24.6.1968. Jugo-
slavia, Crena Gora (Montenegro), Juha Suominen 578, (CAI),
20.6.1971.
Phleum subulatum (Savi) Asch. & Graebn. subsp. ciliatum
(Boiss.) Humphries
Russia, Simkovicset & Kladova, (WU), 667, 1853. Russia,
Simkovicset & Kladova, (WU), 668, 1853. Russia, Kaldova,
(WU), 314, 1853. Russia, Brian Duel, (WU), 1763, 1917. Rus-
sia, Kalmetare, (WU), 976, 1853. Russia, Vlademiaer, (WU),
1867. Russia, (WU), 284, 1889. Russia, Gardeun, (WU),
96721, 1932. Russia, (WU), 121, 1923.
Phleum subulatum (Savi) Asch. & Graebn.subsp. Subulatum
Greece, Karakitsos & Turland, (MO), 5772277, 2003.
Greece, (MO), 1740531, 1955. Palestine, N. Feinbrun & M.
Zohary, (MO), 2631470, 1928. Palestine, N. Feinbrun & M.
Zohary, (WU), 233, 1928. J. Prudhomme, (WU), 4955, 1984.
Poland, A. Schuttz, (MO), 3048084, no date. Greece, Handel-
Mazzetti, (WU), 37, 1907. Poland, M. Gandoger, (MO),
761104, 1914. Syria, (MO), 3048087, 1856. France, A. Bues,
(MO), 3048077. Australia, M. Gandoger, (MO), 763529, 1914.
Palestine, Arjch Obraham, (MO), 1702285, June 1953. Italy, T.
B. Wolfe Keck, (MO), 1822063, 1854.
Webbia: Journal of Plant Taxonomy and Geography 35
