Taxonomic study on the genus Eremopyrum (Ledeb.) Jaub. et Spach (Poaceae) in Turkey

ORIGINAL ARTICLE

Taxonomic study on the genus Eremopyrum (Ledeb.) Jaub. etSpach (Poaceae) in Turkey

Evren Cabi • Musa Dogan

Received: 19 January 2009 / Accepted: 12 May 2010 / Published online: 25 June 2010

� Springer-Verlag 2010

Abstract In this study, we examined whether a satisfactory

infrageneric grouping in Eremopyrum (Ledeb.) Jaub. &

Spach can be achieved with the help of cluster analysis based

on external vegetative and floral morphological characters of

specimens either gathered from the field or borrowed from

various herbaria. Twenty morphological characters were

chosen and scored for 36 populations representing the genus.

The data were subjected to numerical taxonomic analyses,

and a phenogram was obtained by applying Gower general

similarity index. The results of this study suggest that there

are basically five species in Turkey, namely E. triticeum

(Gaertner) Nevski, E. orientale (L.) Jaub. & Spach,

E. distans (K. Koch) Nevski, E. bonaepartis (Sprengel)

Nevski, and E. confusum Melderis. For the first time a new

variety, E. bonaepartis (Sprengel) Nevski. var. aristatum

Cabi & Dogan, is proposed along with two new combinations:

E. bonaepartis (Sprengel) Nevski. var. sinaicum (Steudel)

Cabi & Dogan and E. confusum Melderis var. sublanugi-

nosum (Drob.) Cabi & Dogan. An account of Eremopyrum in

Turkey is given, including a key to the species and varieties,

and a general indication of their distribution.

Keywords Eremopyrum � Poaceae �Multivariate analysis � Turkey

Introduction

Eremopyrum (Ledeb.) Jaub. & Spach is a well-circum-

scribed genus with annual habit and oblong to orbicular

fragile spike-like inflorescence that is used in wheat

improvement (Clayton and Renvoize 1986). It was first

recognized as a taxon by Ledebour (1853), who treated it

as Triticum sect. Eremopyrum. Bentham and Hooker

(1883) transferred it to Agropyron Gaertn., leaving it as a

section. It was Jaubert and Spach (1851) who first recog-

nized Eremopyrum as a genus, distinguishing it from

Agropyron on the basis of its annual habit. Since then 18

species and many infraspecific taxa have been described,

but the number of species currently accepted varies from

four (Tzvelev 1976; Sakamoto 1979; Frederiksen 1991) to

nine (Love 1984).

This genus includes both diploid and tetraploid taxa.

The diploid species E. triticeum (Gaertn.) Nevski and

E. distans (K. Koch) Nevski are well circumscribed and

more easily distinguished than the other species. E. ori-

entale (L.) Jaub. & Spach is a tetraploid, thought to be

derived from the diploid parents E. triticeum and E. distans

via allopolyploidization (Sakamoto 1979). E. bonaepartis

(Spreng.) Nevski, a species complex, includes both diploid

(2n = 14) and tetraploid taxa (2n = 28) (Melderis 1985).

Sakamoto (1979) proposed that tetraploid taxa in

E. bonaepartis originated from the diploid parents

E. bonaepartis and E. distans. Melderis (1985) indicated

that characters which can be used to separate diploid and

tetraploid taxa include the presence or absence of indu-

mentum on the spikelets and the length of the lemma awn.

Eremopyrum consists of the species with F genome

(Love 1984). Intergeneric hybridizations including Ere-

mopyrum show that there are strong sterility barriers

between Eremopyrum and the species included in crossing

efforts (Sakamoto 1967, 1968, 1972, 1974; Frederiksen and

Bothmer 1989; Frederiksen 1993, 1994).

Bor (1968) recognized four species of Eremopyrum in Iraq,

including E. confusum Melderis, which supposedly differs

E. Cabi (&) � M. Dogan

Department of Biological Sciences, Faculty of Arts and

Sciences, Middle East Technical University, Ankara, Turkey

e-mail: [email protected]; [email protected]

123

Plant Syst Evol (2010) 287:129–140

DOI 10.1007/s00606-010-0306-1

from E. bonaepartis in having glumes and lemmas rather than

awnless, sharp pointed glumes and lemmas. In Iran, Bor

(1970) accepted the presence of four species. In Flora Euro-

pea, Melderis (1980) recognized three species, E. triticeum,

E. orientale, and E. distans, in the genus Eremopyrum.

In the Flora of Turkey, Melderis (1985) recognized four

species and two subspecies, namely E. distans, E. orien-

tale, E. triticeum, E. bonaepartis subsp. bonaepartis (Spr-

eng.) Nevski, and E. bonaepartis subsp. hirsutum (Bertol.)

Melderis. According to Melderis, E. bonaepartis covered

diploid (2n = 14) and tetraploid taxa (2n = 28) which

showed huge variability regarding their spikelet characters,

such as size of spikelets, indumentum, shape of glumes,

and length of lemma awn. He also indicated that Sprengel’s

material was obtained from cultivated material from Egypt.

He treated diploid and tetraploid taxa as different subspe-

cies in Turkey (Melderis 1985). After publication of the

flora, the new grass taxa published were compiled in the

supplements (Davis et al. 1988; Guner et al. 2000). In

addition to these, taxonomical and palynological studies

were performed on certain genera (Dogan 1988, 1991,

1992, 1997; Cabi and Dogan 2009; Cabi et al. 2009; Ozler

et al. 2009; Cabi et al. 2010).

The most recent revisional study on the genus was con-

ducted by Frederiksen (1991). In her study, she recognized

four species, namely E. triticeum, E. orientale, E. distans,

and E. bonaepartis. She stated that E. bonaepartis included

three morphologically distinct subgroups previously

accepted as separate species by different authors: E. con-

fusum, characterized by having distinctly awned glumes and

lemmas; E. bonaepartis s. str., with sharp pointed glumes

and lemmas; and Triticum sinaicum Steud., characterized by

gradually tapering glumes and lower 1–3 lemmas, but dis-

tinctly awned upper lemmas. She placed these three mor-

phologically different groups under the same species, since

there was not a clear-cut delimitation among the examined

groups. In her study, she used seven characters based on

glume and lemma morphology to separate the taxa included

in E. bonaepartis. She omitted the indumentum characters

because they seemed to be difficult to quantify. She also

constructed a scatter diagram based on dimensions of

glumes and lemmas and presence of glume and lemma awn.

This scatter diagram showed that there were no significant

differences among these divergent groups.

Frederiksen (1991) also made some chromosome counts

and noted that Eremopyrum confusum populations were

found to be tetraploid. Typical E. bonaepartis s. str. was

predominantly tetraploid, while Triticum sinaicum was

always diploid.

The aim of this study is to determine the morphological

variation pattern in this genus via numerical analyses and

also outline the taxonomic status of the infrageneric

grouping in the genus in Turkey.

Materials and methods

Specimens reviewed in this assessment were either gathered

from their natural habitats or borrowed from major herbaria

such as ANK, GAZI, ISTE, ISTF, VANF, K, E, and BM

(herbarium abbreviations according to Holmgren et al.

1990). Most of the studied material was collected from

different localities of Turkey during field surveys carried

out as part of a research project sponsored by the Turkish

Scientific and Technical Research Council (TUBITAK) for

a 3-year period starting from 2006. The material was

deposited in the Plant Systematics and Biodiversity Labo-

ratory in Biological Sciences Department of Middle East

Technical University. Identification of specimens was

achieved by using keys given in the relevant studies by Bor

(1968), Melderis (1985), and Frederiksen (1991).

Some of these herbarium specimens and collected

specimens were selected for multivariate analysis as

operational taxonomic units (OTUs). A list of OTUs for

numerical analysis is given in Appendix I. Twenty diag-

nostic characters (Tables 1, 2) were chosen and scored for

each OTU (Sneath and Sokal 1973). Morphological mea-

surements on each OTU were made using a Leica L2 ste-

reomicroscope and Leica Application Suite software

package. The raw data matrix includes quantitative and

ordinal variables. For the purpose of morphometric anal-

ysis, Gower’s (1971) general similarity index was chosen

to generate a distance matrix. This distance matrix was

used for cluster analysis with the help of the Unweighted

Pair Group Method with Arithmetic mean (UPGMA)

algorithm. Cluster analysis as implemented in MVSP

software version 3.1 (Kovach 1999) was performed by

using the same distance matrix. The advantage of Gower’s

coefficient is that it allows the presence of a mixture of all

variable types and tolerates missing values as well (Mason

et al. 2005).

Table 1 Quantitative characters studied in Eremopyrum genus

No. Abbreviation Character

1 SL Spike length (cm)

2 SW Spike width (cm)

3 FPS Florets number per spikelet

4 SptL Spikelet length (mm)

5 GL Glume length (mm)

6 GW Glume width (mm)

7 GAL Glume awn length (mm)

8 LeL Lemma length (mm)

9 LeAL Lemma awn length (mm)

130 E. Cabi, M. Dogan

123

Results and discussion

Quantitative characters (Table 1) used to construct the

phenogram showed large variation (Fig. 1). There seems to

be no single diagnostic character that can be used for

recognition of species, but only a combination of these

characters can be useful. Lengths of glumes’ and lemmas’

awns varied among the representatives of the same popu-

lations collected at different times. Its small spike distin-

guished E. triticeum from the remaining taxa found in this

genus. The narrowest glume size was found to be present in

E. distans populations. Glumes are distinctly shorter than

lemmas in diploid E. bonaepartis (=T. sinaicum) popula-

tions compared with E. confusum or E. bonaepartis subsp.

hirsutum populations.

Among the studied qualitative characters (Table 2)

rachis fragility, shape of glumes, frequency of hairs on

spikelet, glume and lemma, keels of palea having two short

awns with or without a deep cleft were found to be the most

important qualitative diagnostic characters (Table 3).

The phenogram (Fig. 2) shows two major clusters, repre-

senting the sections used by Nevski (1936). Both clusters

separate at \0.6 similarity level. The first main cluster

includes E. triticeum populations, which have tough rachis

(sect. Micropyrum), and the second cluster includes the

remaining taxa, which have fragile rachis (sect. Eremopyrum).

Cluster 2 includes E. bonaepartis populations with

sharply pointed glumes and lower lemmas (the lowest 1–2

lemmas), while upper (1–2) lemmas have a distinct awn.

OTU 20 is separated because of its awn of glumes and

lemmas. This taxon has an up to 1.5-mm-long glume awn,

and its lowest lemma has a distinct awn up to 2.5 mm long.

However, this taxon has been treated as a new variety,

E. bonaepartis (Sprengel) Nevski var. aristatum Cabi &

Dogan. It is found that there is a gradual increase in length of

awns of lemmas from the lowermost lemma to the upper-

most one. When the lowest lemma is awnless, the upper

lemmas usually have a short awn but absolutely bigger than

former lemma awn. Although typical E. bonaepartis popu-

lations have sharp pointed glumes and lemmas are observed

in the field in the early stage, it is determined that awns are

formed on the upper one or two lemmas in the later stages.

Cluster 3 was composed of the populations of

E. distans, which is one of the clearly circumscribed

species found in this genus. It differs from other species in

having keels of palea prolonged into two short awns

(0.7–2 mm) with a deep cleft between them. It also differs

in having narrowly linear to lanceolate glumes equal in

length to the lemmas or slightly longer.

Cluster 4 includes E. orientale populations, which are

characterized by strongly curved glumes and lemmas. The

pubescence of the glumes is among the most variable

character in this cluster. In some populations of E. orientale

(OTU 6 and 7) the glumes are very sparsely hairy or gla-

brous, which is why they form a subcluster under cluster 4.

Owing to the close relationship within the populations in

cluster 5, we observed three main subclusters in this clus-

ter. Cluster 5 was composed of populations belonging to

E. bonaepartis subsp. hirsutum and E. confusum. The

common characteristics of these populations are as follows:

All populations have long glumes relative to E. bonae-

partis populations. Glumes are slightly shorter than

lemmas or equal in length to lemmas. There was no gradual

change in the awns of lemmas found in the E. bonaepartis

populations. Differences are observed in the indumentum

of spikelets and the awn length of glumes and lemmas.

E. confusum has distinctly awned glumes and lemmas,

whereas E. bonaepartis subsp. hirsutum populations have

gradually tapering glumes and lemmas.

The results of this study confirmed that the diploid

E. bonaepartis (=T. sinaicum) is morphologically distinct

and recognizable. Its members have sharp pointed or shortly

awned glumes and lower 1–2 lemmas, and distinctly awned

upper lemmas. The awns of lemmas are characterized by

gradual increase in length from the lowermost lemmas to the

upper ones. In all populations of E. bonaepartis collected

from Turkey we also observed morphological variation

regarding awn length of lemma, but there is always a gradual

increase from the lowest lemma to the uppermost one. This

feature is not seen in any other taxa, such as E. bonaepartis

subsp. hirsutum and E. confusum. The indumentum of

glumes and lemmas of E. bonaepartis is generally scabrid to

dense scabrid especially on the keel, but in early stage this

taxon has glabrous glumes and lemmas; E. bonaepartis

usually has distinctly shorter glumes than lemmas.

In contrast to Frederiksen (1991), our results indicated

clear differences among the subgroups found in

Table 2 Qualitative characters studied in Eremopyrum

No. Abbreviation Character

1 SpHT Spike hair type (glabrous 0, scabrous 1, hairy 2)

2 GS Glume shape (lanceolate 0, angustiform 1, curved

2)

3 GH Glume hair (smooth 0, scabrid 1, villous 2)

4 FGH Frequency of glume hair (dense 0, sparse 1)

5 SGA Shape of glume apex (awned 0, unawned 1)

6 Paapx Palea apex (gapped 1, no gapped 0)

7 Rchf Rachis fragility (fragile 0, not fragile 1)

8 Rchlf Rachilla fragility (fragile 0, not fragile 1)

9 GvL Glume length versus lemma length (glumes equal

or subequal to lemmas 0, glume distinctly

shorter than lemmas 1)

10 AoL Awn of lemmas (unequal 0, subequal or equal 1)

11 AoLL Apex of lowest lemma (with a mucro 0–1 mm

long 0, with a distinct long awn 1)

Taxonomic study on the genus Eremopyrum (Ledeb.) Jaub. et Spach (Poaceae) 131

123

E. bonaepartis complex. She constructed the scatter diagram

by using only three morphological characters: glume length,

lemma length, and existence of glume and lemma awn.

Although she noted that Triticum sinaicum possesses an awn

on the upper lemmas, she did not record this character in

T. sinaicum populations while constructing the scatter dia-

gram. She took into account only the lowest lemma charac-

teristics. Although she realized that T. sinaicum was rather

distinct from typical E. bonaepartis and E. confusum as the

glumes and lower 1–3 lemmas were blunt while the upper

lemmas had a distinct awn, she did not use these differences

in awn length of lemmas within a spikelet.

Taking all the above-mentioned facts into consideration,

we recognized five species including the E. confusum for

Flora of Turkey.

Taxonomic treatment

Eremopyrum (Ledeb.) Jaub. & Spach

III. Pl. Or. 4: 26 (1851) p.p.—Triticum L. sect. Eremopy-

rum Ledeb., Fl. Alt. 1: 112 (1829) p.p.—Agropyron

Gaertner sect. pycnopyrum K. Koch, Linnea 21: 425(1848)

p.p.—Type: E. orientale (L.) Jaub. & Spach [lectotype,

selected by Nevski, Trudy Bot. Inst. Akad. Nauk SSSR,

ser. 1,2: 63 (1936)].

Annuals. Stem fasciculate or geniculate. Leaf blades

linear acuminate, flat, folded when dried; auriculate, auri-

cles very small or inconspicuous. Inflorescence a dense

terminal spike; rachis tough or fragile, with short inter-

nodes. Spikelets solitary, laterally compressed, alternately

and distichously arranged at rachis nodes, widely diver-

gent, with 2–6 florets, upper florets ±rudimentary. Glumes

boat-shaped, coriaceous, becoming indurate at maturity,

with prominent asymmetric keel, gradually tapering to a

±sharp point or more or less distinct awn, the edges of

glume with broad hyaline margins. Lemma coriaceous,

rounded on back, keeled above, five veined, unawned or

awned with a more or less distinct awn. Palea 2-keeled

with more or less conspicuously cleft between keels.

Anthers yellow, 0.4–1.4 mm long. Caryopsis adherent to

palea at maturity.

Eremopyrum (Ledeb.) Jaub. & Spach

Key to the species and varieties of Eremopyrum recognized

in this study

Fig. 1 Box plot of quantitative

characters screened for cluster

analysis. SL spike length (cm),

SW spike width (cm), FPSflorets number per spikelet, SptLspikelet length (mm), GL glume

length (mm), GW glume width

(mm), GAL glume awn length

(mm), LeL lemma length (mm),

LeaL lemma awn length (mm)

Table 3 Morphological differences between E. bonaepartis and

E. confusum species

E. bonaepartis E. confusum

Spikelets 8–15.5 mm long,

imbricate

Spikelets 10–20 mm long,

spreading

Spikelets glabrous or more or less

scabrid

Generally densely covered with

long hairs or sometimes

glabrous

Glumes 6–11 mm long, gradually

tapering to a short awn or

sharply pointed, distinctly

shorter than lemmas

Glumes 9–16 mm long, slightly

shorter or equal to lemmas

Upper lemmas have awns longer

than lower lemmas

The length of lemma awns equal

or subequal, there is no gradual

change in length


123

1. E. triticeum (Gaertner) Nevski in Acta Univ. Asiae

Med. ser. 8b (Bot.) 17:52 (1934).

Syn: Agropyron triticeum Gaertner in Novi Comment.

Acad. Sci. Petrop. 14:540, t. 19 f. 4–5 (1770); Secale

prostratum Pall., Reise Abh. 1:485 (1771); Triticum pro-

stratum (Pall.) L. f., Suppl. Pl. 114 (1781); Agropyron

prostratum (Pall.) P. Beauv., Ess. Agrostogr. 102 & 146

(1812). Ic: Fl. URSS 2: t. 47a (‘‘67a’’) f. 7 (1934);

Sakamoto in Mem. Coll. Agr. Kyoto Univ. 114:9, f. 1i, j,

11 f. 3, i, j (1979)—Type: Pallas, in desertis caspiis [LINN

104,18 lectotype selected by Frederiksen (1991)].

Stems 4–30 cm, geniculate at base, scabrid or retrorsely

pilose towards spike. Leaf blades 1.4–5 9 0.15–0.4 cm,

scabrid or shortly pilose above; sheaths partly cover the

spike. Spike 0.9–1.7 cm, elliptic-ovate or nearly orbicular

in outline. Spikelets 6–10 mm, usually with 3–4 florets;

rachilla fragile, except lowermost. Glumes 4–6.5 mm,

broadly lanceolate, prominently inflated at base, strongly

curved, glabrous and subulate at apex. Lemma 5–7.5 mm

with an awn up to 2 mm, keel prominent on upper half of

lemma. Palea 3.5–4 mm with a low cleft at apex, slightly

shorter than body of lemma.

Fl. 5–6. Steppe, sandy arid places, Alt. 850–1,715 m.

This species grows sympatrically with other Eremopyrum

species such as E. orientale and E. distans.

Described from the valley of the River Yaika (in the

Ural Mts., Russia).

North and Inner Anatolia (Fig. 3). A7 Gumushane:

Bourgeau. A9 Kars: ruins of Ani, 27 vi 1967, J.G. Ross;

B2 Usak: Usak, 915 m, Bal. 1857:1339; B4 Konya:

Yavsan Memlehanesi nr Tuz Lake, 908 m, E. Cabi 1706;

Ankara: 15 km W. of Ankara, 900 m, Ehrend. 62-1/79-1;

B5 Yozgat: Yozgat to Bogazlıyan, Kaskısla Village,

1682 m, E. Cabi 3274; B7 Erzincan: Egin (Kemaliye),

Sint 2370!; B8 Erzurum: nr Erzurum, vi 1853, Huet (with

E. distans)!; B9 Erzurum: Horasan Town centre, 1,715 m,

E. Cabi 2492; Van: nr Van, Sakamoto (Kyoto Univ. Bot.

Expend.); Van Canik Tuzlası, 1,750 m, L. Behcet 4597!;

Key to the species and varieties of Eremopyrum recognized in this study

1. Rachis tough, not breaking off at maturity; spikes up to 1.7 cm. 1. E. triticeum

1. Rachis fragile, breaking off at maturity; spikes longer than 1.5 cm.

2. Glumes strongly curved 3. E. orientale

2. Glumes not curved or slightly curved at maturity

3. Paleas with terminal two short awns (0.7-2 mm) with deep cleft between them

4. E. distans

3. Paleas with two very short teeth (up to 0.4 mm) with shallow or moderately cleft

between them.

4. Spikelets imbricate; glumes distinctly shorter than lemmas, there is gradual

increase in the length of awn of lemmas 2. E. bonaepartis

5. Glumes and lowest lemma gradually tapering into a sharp point

2a. E. b. var. sinaicum

5. Glumes and lowest lemma have a distinct awn

2b. E. b. var. aristatum

4. Spikelets spreading; glumes subequal or equaling to lemmas, awn of lemmas are

subequal, there is not any gradual change 5. E. confusum

6. Glumes and lemmas gradually tapering into a sharp point

5a. E. c. var. sublanuginosum

6. Glumes and lemmas distinctly awned

7. Spikelets glabrous 5b. E. c. var. glabrum

7. Spikelets covered densely with more or less long white hairs or with

short rigid bristles 5c. E. c. var. confusum


123

B10 Kars: 27 km from Igdır to Aralık (Aras Valley),

880 m, D. 43654!; C3 Burdur: Burdur, Heldr. C4 Konya:

Kucuk Koy, 980 m, Helbaek 2423!

S.E. Europe, Crimea, Caucasia, Iran, C. Asia, Siberia.

2. E. bonaepartis (Sprengel) Nevski in Acta Univ.

Asiae Med. ser. 8b (Bot.) 17:52 (1934).

Syn: Triticum bonaepartis Sprengel, Erst. Nachtr. Bot.

Gart. Halle 40 (1801); T. squarrosum Roth, Neue Beytr.

Bot. 1: 128 (1802); Eremopyrum squarrosum (Roth) Jaub.

& Spach, Ill. Pl. Or. 4: t. 320 f. 2 (1851); Agropyron

kotschyanum Boiss. & Hohen. in Boiss., Diagn. ser. 1(13);

69 (1854)! Ic: Fl. URSS 2: t. 47a (‘‘67a’’) f. 8a–c (1934), as

E. buonapartis; Fl. Iraq 9:t. 78 f. 2 (1968), as var.

bonaepartis; Sakamoto, op. cit. 9, f. 3a (1979). 2n = 14—

Type: Schimper 157, Bastan ad radices montis Sinai, 1835

[P ex herb. Steudel, lectotype, selected by Frederiksen

(1991)], G, G-herb. Boissier, K, M, P, TUB paralectotypes.

Stem 1.5–32.5 cm, geniculate at base, glabrous but

scabrid below either spike and nodes, 2–3 noded. Leaf

blades 2–10 9 0.2–0.4 cm, flat to folded, convolute when

dried, scabrid above; uppermost sheath slightly inflated,

partly enclosing the spike. Spike 1.3–4 9 1.1–1.5 cm,

oblong, elliptic to ovate, obtuse or truncate at apex. Spik-

elets 8.5–15 mm with 3–4 florets, the lower 2–3 florets

Fig. 2 Phenogram constructed by means of Gower general similarity index. Cl cluster number; OTUs operational taxonomic units

Fig. 3 Distribution of

Eremopyrum triticeum (filledsquares) in Turkey


123

fertile, the upper one rudimentary; rachis fragile, rachilla

tough. Glumes strongly keeled asymmetrically, more or

less scabrid on keel, lanceolate, distinctly shorter than

lemmas, obtuse or acuminate at apex. Lemma 8.5–15 mm

(incl. awn), glabrous or scabrid, linear lanceolate, keeled

rounded at the back, awn 0–6 mm sometimes the lowest

lemmas with a mucro (c. 0.5–1 mm) sparsely scabrid all

over at the back, upper ones with an awn (2–6 mm, but

generally the lowest lemma has shorter awn than upper

ones). Palea 4.5–5.5 mm, distinctly shorter than lemmas,

apex with a short cleft, scabrid on keels.

Var. sinaicum (Steudel) Cabi & Dogan, comb. nov.

Syn: Triticum sinaicum Steudel Syn. Pl. Glum. 1: 346

(1855); Dasypyrum sinaicum (Steudel) Candargy, Arch.

Biol. Veg. Pure Appl. 1: 35, 62 (1901)—Type: Schimper

157, Bastan ad radices montis Sinai, 1835 [P ex herb.

Steudel lectotype, selected by Frederiksen (1991)], G,

G-herb. Boissier, K, M, P, TUB paralectotypes.

Fl.4–6 Hab. Salty dry steppes, slopes, roadsides, field

margins. Alt. 454–1,200 m. Grows with Aegilops cylind-

rica Host, Ae. speltoides Tausch, Hordeum murinum L.,

Leymus cappadocicus (Boiss. & Balansa) Melderis,

E. orientale, and E. confusum.

Inner, South, and East Anatolia (Fig. 4). B3

Eskisehir: 29 km from Polatlı to Sivrihisar, 800 m, Coode

and Jones 2265 (with E. confusum)!; B4 Ankara: Ankara

to Sereflikochisar, 12 km before Sereflikochisar, shore of

Tuz Lake, 912 m. E. Cabi 3059; Konya: Cihanbeyli Tuz

enterprise, shore of Tuz Lake, 909 m. E. Cabi 2246 (with

E. orientale and E. confusum); B7 Malatya: nr Gozene,

1970, Sakamato (Kyoto Univ. Bot. Exped.); B8 Erzurum:

Kassuklu Valley between Bayburt and Erzurum, v 1853,

Huet!; B10 Kars: nr Aralık (Grossheim 1: map 312); C3

Antalya: 36 km from Elmalı to Korkuteli, Alava 5257; C5

Konya: Eregli to Ulukısla, 1,200 m, It. Leyd. 1959:1014

(with E. confusum)!; C7 Sanlıurfa: Ceylanpınar State

Farm, Seyh Nasr streambed, 454 m, E. Cabi 2241; S.Urfa:

S.Urfa to Akcakale, 13 km from S.Urfa, 500 m, D. 28141!;

C8 Mardin: 10 km from Gercus to Hasankeyf, 792 m, E.

Cabi 2254; Diyarbakır: Diyarbakır, nr Kupri, Kotschy

1841:142.

N. Africa, Iran, Syrian Desert, Sinai. Ir.-Tur. element.

Var. aristatum Cabi & Dogan var. nova (Fig. 5a)

Var. sinaicum smilis, ab ea diversa: glumae ab infima

lemmatum vel aristata

Type: Turkey B3 Ankara: 26 km from Polatlı to Siv-

rihisar 39�33.4170N 31�47.1730E 814 m steppe, E. Cabi

339 (holotype: GAZI; isotypes ANK, E).

Fl 4–5. Hab. Steppe, roadsides, gypsum fields. Alt.

810–1,140 m

Etymology: The name derives from its aristate lowest

lemma

Central Anatolia (Fig. 4). B4 Konya: Kulu to Konya c.

7–8 km before Konya, 39�01.1490N 33�01.7440E, road-

sides, steppe, 1,136 m, E. Cabi 2243; B6 Malatya: 1 km


Eremopyrum bonaepartispopulations with two varieties,

var. aristatum (open squares)

and var. sinaicum (filledsquares), in Turkey

Fig. 5 Spikelets of a Eremopyrum bonaepartis var. aristatum,

b Eremopyrum confusum var. confusum


123

from Darende to Malatya, 38�3003200N 37�3102400E1,011 m, roadsides, gypsum fields A.K.1103!

3. E. orientale (L.) Jaub. & Spach, III. Pl. Or. 4:26, t.

319 (1851).

Syn: Secale orientale L., Sp. Pl. 84 (1753); Agropyron

orientale (L.) Roem. & Schult., Syst. Veg. 2:757 (1817);

A. bourgeaei Boiss., Fl. Or. 5:669 (1884)! Ic: Fl. URSS 2:

t. 47a (‘‘67a’’) f. 9 a–d (1934); Fl. Iraq 9: t. 81 (1968);

Sakamoto in Mem. Coll. Agr. Kyoto Univ. 114:9, f. lh, 11,

f. 3 h(1979).

Fl. 3–5 Hab. Steppe, lake margins, canal banks, hill-

sides, usually on clayey and muddy ground. Alt. 980–

1,825 m.

Stems 4–17 cm. Leaf blades 2–3 mm broad, ±scabrid

on both surfaces, sometimes shortly pilose above. Spike to

2.5 cm, usually ovate-elliptic, long-exserted. Spikelets

10–15 mm, pilose, with 2–3 florets; rachilla tough. Glumes

5–6.5 mm (excl. awn), more or less covered by long white

hairs, lanceolate, strongly curved, prominently veined,

gradually tapering to a short awn (2–5 mm). Lemma as

long as glumes, with prominent keel and short awn

(2–4 mm). Keels of palea prolonged into two short teeth

(0.4–0.6 mm), with deep cleft between them.

Described from islands of the Aegean Sea.

Central, North, and East Anatolia (Fig. 6). A5

Amasya: Merzifon to Kamıslı, A. Baytop (ISTE 9124)!;

A7 Trabzon/Erzurum: ‘Armenia’, Calvert & Zohrab

(with Agropyron incanum (Nabelek) N.N. Tzvelev); A8

Erzurum: W. of Ispir, 1,400 m, Barclay 784!;

Gumushane: Bayburt, Bourgeau (type of Agropyron

bourgeaei)!; A9 Kars: Kagızman (Grossheim 1:

map 313); B4 Ankara: 12 km to Sereflikochisar, nr Tuz

Lake., 912 m, E. Cabi 3055; Konya: inside of Cihanbeyli

Salt enterprise area, shore of Tuz Lake, 909 m, E. Cabi

2244a (with E. bonaepartis and E. confusum); Konya:

Yavsan, nr Tuz Lake, D. 18697!; B5 Yozgat: Yozgat to

Bogazlıyan, Kaskısla Village, 1682 m, E. Cabi 3273;

Kayseri: Karapınar, 1,000 m, 1902, Penther &

Zederbauer!; B7 Elazıg: Baskil, Karabas Village, 1500 m,

L. Behcet 5211!. B8 Erzurum: Erzurum to Pazaryolu-_Ispir, 1 km before Eskipolat Village 1,825 m, E. Cabi

3429; B9 Agrı: Agrı to Dogubeyazıt, 21 km before Diy-

adin, 1,805 m. E. Cabi 2501; Bl0 Kars: 3–5 km E. of

Aralik (Aras Valley), 850 m, D. 43671!; Van: Ozalp, N.

of Damlacık Village 2,425 m, F. Ozgokce 613!; C4

Konya: 1.5 km N.W. of Alemdar Village, 1,008 m, E.

Cabi 631. Konya: Kucuk Koy, 980 m, Helbaek 2462!; C5

Konya: Pertek, nr Nigde, 1,300 m, Siehe 1912:580!.

N. Africa, S. Russia, Crimea, Caucasia, Iran, Iraq, W.

Syria, Arabia, Afghanistan, C. Asia, W. Pakistan, N.W.

India, Siberia. Ir.-Tur. element.

4. E. distans (K. Koch) Nevski in Acta Univ. Asiae

Med. ser. 8b (Bot.) 17:52 (1934).

Syn: Agropyron distans K. Koch in Linnaea 21 (4):426

(1848); A. lasianthum Boiss., Diagn. ser. 1 (13): 68 (1853);

A. orientale (L.) Roemer & Schultes var. lasianthum

(Boiss.) Boiss., Fl. Or. 5:668 (1884)! Ic: Fl. URSS 2: t. 47a

(‘‘67a’’) f. 6a–b(1934); Fl. Iraq 9: t. 80 (1968). Eremopy-

rum orientale var. lanuginosum (Griseb.) Candargy, Arch.

Biol. Veg. Pure Appl. 1: 59 (1901). Triticum orientale var.

lanuginosum (Griseb.) in Ledeb., Fl. Ross: 4:337 (1853)!—

Agropyron orientale var. lanuginosum (Griseb.) K. Richt.,

Pl. Eur. 1: 126 1890 selected by Frederiksen as isolectotype

K!

Fl. 5-8 Hab. Sandy gravel plains, steppe, igneous

slopes, red soils. Alt. 850–2,500 m.

Type: [Soviet Armenia, Erevan] aus der Provinz Eriwan,

915 m, auf basaltisch-trachytischem Boden, [K. Koch]

(holo. B, destroyed).

Stems up to 15 cm. Leaf blades 2–3 mm broad, scabrid

or sparsely pilose on upper surface. Spike 2.5–5 9 1.5–

2.5 cm, oblong, elliptic oblong. Spikelets 15–17 mm,


Eremopyrum orientale (filledsquares) in Turkey


123

densely villous, with 3–5 florets, the lower florets her-

maphrodite, the upper more or less reduced; rachilla tough.

Glumes 11.5–16 mm (incl. awn), narrowly linear-lanceo-

late, gradually tapering to a short awn (4–7 mm), densely

villous on dorsal side but scabrid towards apex. Lemmas

7.5–13 mm (incl. awn) as long as or shorter than glumes,

elliptic-acuminate, with prominent keel and short awn (4–

7 mm), body of upper lemmas wider than lower ones.

Palea protruding during flowering, keels of palea prolonged

into two short awns (0.7–2 mm), with deep cleft between

them, palea as long as body of lemma.

N.E. and Inner Anatolia (Fig. 7). A8 Gumushane:

Bayburt, Bourgeau 254 (with E. orientale)!; A8/9 Erzu-

rum: Oltu (Grossheim 1: map 314); A9 Kars: Igdır to

Tuzluca (Aras Valley), 920 m, D. 43567b!; Erzurum: Ka-

rakilise (Grossheim 1: map 314); B4 Konya: Yavsan

memlehanesi nr Tuz Lake., Andrasovszky 911!; B5 Konya:

nr Eregli, 18 v 1902, Zederbauer. B8 Erzurum: nr Erzurum,

vi 1853, Huet (with E. triticeum); B9 Van: 7 km from Van to

Ercek, 1850 m, D. 44251!; Van: Gurpınar, Kocguden Vil-

lage, 2,100 m. M. Unal 5356!; Bl0 Kars: 3–5 km from

Aralık (Aras Valley), 850 m, D. 43671!; Agrı: 17 km N. of

Dogubeyazıt towards Igdır, 1600 m, Sorger & Buchner

82-62-45!;. Van: near Ozalp, Sakamoto (Kyoto Univ. Bot.

Exped., 1970); C9 Hakkari: 80–86 km S. of Gurpınar,

Yalınca to Hakkari, 2500 m, Sorger & Kit-Tan 84-78-19!.

S.E. Russia, Caucasia, Iran, Iraq, Afghanistan, C. Asia,

W. Pakistan. Ir.-Tur. element.

5. E. confusum Melderis in Anz. Math.-Nat. Oesterr.

Akad. Wiss., 15 (1964); Bor in Rech. f., Fl. Iraq: 119

(1964); Rawi in Dep. Agr. Iraq Tech. Bull. 14: 207 (1964)

Syn: Eremopyrum bonaepartis subsp. hirsutum (Bertol.)

Melderis in Notes R.B.G. Edinb. 42:81 (1984); Hordeum

hirsutum Bertol., Misc. Bot. 1:11 (1842)!; Eremopyrum

squarrosum (Roth) Jaub. & Spach var. hirsutum Cand. in

Arch. Biol. Veg. Athenes 1:33 (1901)! Agropyron orientale

(L.) Roemer & Schultes var. sublanuginosum Drobov in

Trv. Mus. Bot. Acad. Sci. Petersb. 16:135 (1916); Ere-

mopyrum hirsutum (Bertol.) Nevski in Acta Univ. Asiae

Med. ser. 8b (Bot.) 17:52 (1934)! E. bonaepartis (Spren-

gel) Nevski var. sublanuginosum (Drobov) Melderis in

Ark. Bot. ser. 2, 2:305 (1952). Ic: Fl. Iraq 9: t. 7 f. 3–10

(1968).

Stem 1.5–32.5 cm long, geniculate, densely hairy below

spike, 2–3 noded. Leaf blades up to 10 cm long and 4 mm

broad, flat to folded, convolute when dried, scabrid above;

sheaths partly enclose the spike. Spike 2.7–4 9 1–2.3 cm,

oblong, elliptic, obtuse or truncate at apex. Spikelets cov-

ered with long white hairs or glabrous with 4–6 florets, the

lower 3–5 florets fertile, the upper one rudimentary; ra-

chilla tough. Glumes 10.5–14.5 mm, curved slightly

towards apex, narrowly lanceolate, keeled asymmetrically,

slightly shorter than or equal to lemmas, usually gradually

tapering or distinctly awned at apex with an unconspicuous

awn (0.5–1.5 mm). Lemma linear lanceolate, 13.5–15 mm

(incl. awn), awn of lemma up to 2 mm long. Palea dis-

tinctly shorter than lemmas, 6.5–8 mm, apex of palea with

a short cleft, scabrid on keels.

Described from Euphrates (Upper Euphrates), Chesney

Exped. 196 (holo. BOLO, iso. K!).

Var. sublanuginosum (Drob.) Cabi & Dogan, comb.

nov.

Syn: Agropyron orientale var. sublanuginosum Drob. in

Trav. Mus. Bot. Acad. Sc. Petersb., 16: 135 (1916);

E. bonaepartis var. sublanuginosum (Drob.) Melderis in

Ark. Bot. ser. 2, 2: 305 (1952); Eremopyrum bonaepartis

subsp. hirsutum (Bertol.) Melderis in Notes R.B.G. Edinb.

42:81 (1984).

Glumes gradually tapering, sharp pointed, covered more

or less hirsute hairs.

Central and East Anatolia (Fig. 8). B4 Aksaray:

Aksaray to Sereflikochisar, around Tuz Lake, 970 m, G.


Eremopyrum distans (filledsquares) in Turkey


123

Akaydın 7764; Konya, Kulu to Konya c. 7–8 km

39�01.1490N 33�01.7440E, roadsides, steppe, 1,136 m, E.

Cabi 2242; B5 Nevsehir: 4 km W. of Goreme 1,130 m,

GAZI 5091; B6 Sivas: Gurun, E. of Gokpınar Lake,

38�39042.500N, 37�18014.600E, 1,515 m, E. Cabi 2440;

Sivas; Bogazlıdere to Kutlukaya, 4 km after Kutlukoy,

39�22.9490N 36�55.7260E, 1,471 m, E. Cabi 1306; Sivas:

Ulas to Kangal, 2 km before Kangal, 39�12.1110N37�14.5370E, 1,534 m, E. Cabi 3632; B7 Elazıg: Hazar

Golu, ANK28969; B9 Agrı: Agrı to Dogubeyazıt, 17 km

before Dogubeyazıt, E. Cabi 2508; B9 Van: Van to Ercis

10–15 km 1,800 m. G. Akaydın 7690; C4 Konya

Cihanbeyli, Tersakan Lake. around Alkim, salty marshes,

1,025 m, G. Akaydın 7724.

Fl. 4–7 Hab. Eroded calcareous slopes, salty marshes.

Alt. 950–1,800 m.

Var. confusum (Fig. 5b)

Glumes and lemmas distinctly awned, covered more or

less with long white hairs or with short rigid bristles.

Central Anatolia (Fig. 8). B4 Ankara: Ankara to

Sereflikochisar, around Tuz Lake, G. Akaydın 7727b; B5

Kayseri: Sultanhanı to Bunyan, 5 km from Sultanhanı,1,203 m, 38�5703900N 35�5301900 E, E. Cabi 2249; B6

Sivas: Ulas to Gurun, junction of Deliilyas and Altınyayla

Villages, 39�23047.200N, 37�04059.300E, 1434 m, limestone

hills, E. Cabi 3383; C4 Konya; Cihanbeyli Tuz _Isletmesi;

near Tuz Lake, E. Cabi 2245.

Fl. 5–6 Hab. Sandy and saline soils, slopes and lime-

stone hills. Alt. 950–1,450 m.

Var. glabrum Melderis in Taxon 16:68 (1967)

East Anatolia (Fig. 8). B10 Agrı: 9.5 km from

Dogubeyazıt to Igdır, 39�39026.800N, 44�02034.600E,

1,545 m. E. Cabi 2255.

Fl. 6–7 Hab. dry volcanic rocks, slopes. Alt. 1,545 m.

Acknowledgments Special thanks to Mary Barkworth for critical

reading of the manuscript. Thanks are due to the curators of herbaria

ANK, GAZI, ISTE, ISTF, VANF, E, K, and BM, who allowed us to

study their Eremopyrum specimens, and to the Scientific and Tech-

nical Research Council of Turkey (TUBITAK-TBAG-105 T 171) for

their financial assistance for the study.

Appendix I: list of specimens used in constructing

the cluster analysis (the number in brackets refer

to the OTU numbers seen on phenogram)

E. confusum

G. Akaydın 7764 Ankara: Ankara to Sereflikochisar,

around Tuz Lake (OTU 21).

E00257932 E. confusum var. pakistanicum. Type of var.

pakistanicum (OTU 22).

E. Cabi 2245 Konya: Cihanbeyli, inside of salt enter-

prise, edge of Tuz Lake (OTU 23)

E00257923 _Icel: 5 miles from Mut to Karaman (OTU

24)

G. Akaydın 7727b Ankara: Ankara to Sereflikochisar,

around Tuz Lake (OTU 25)

E. Cabi 2249 Kayseri: Sultanhanı to Bunyan, 5 km from

Sultanhanı (OTU 26)

E. Cabi 2240 Sivas: Gurun, East of Gokpınar Lake

(OTU 27)

G. Akaydın 7690 Van: Van to Ercis *10–15 km (OTU

28)

G. Akaydın 7724 Konya: Cihanbeyli, around Tersakan

Lake (OTU 29).

G. Akaydın 10318 Ankara: Cayırhan Bird Sanctuary.

(OTU 30)

E. Cabi 1306 Sivas: Bogazlıdere to Kutlukaya

(OTU31)

E. Cabi 2508 Agrı: Agrı to Dogubeyazıt 17 km before

Dogubeyazıt (OTU 32)

ANK 28969 Elazıg: Hazar Lake (OTU 33)


Eremopyrum confusum with

three varieties, var.

sublanuginosum (filled squares),

var. confusum (open squares),

and var. glabrum (asterisk), in

Turkey


123

GAZI 5091 Nevsehir: 4 km W. of Goreme (OTU 34)

E. Cabi 2255 Agrı: 9.5 km from Dogubeyazıt to Igdır,(OTU 36)

E. orientale

GAZI 6124 Ankara: Sincan, Fatih Mh.(OTU 13).

E. Cabi 631 Konya: 1.5 km N.W. of Alemdar Village

(OTU 7).

E. Cabi 8 Konya: Cihanbeyli, around Tuzla, Karatepe

Village (OTU 6).

E. Cabi 2501 Agrı: Agrı to Dogubeyazıt, 21 km before

Diyadin (OTU 10).

E. Cabi 2507 Agrı: Agrı to Dogubeyazıt (OTU 9).

E. Cabi 2493 B9 Erzurum: Horasan Town centre (OTU

8).

E. Cabi 2244 Konya: Cihanbeyli, shore of Tuz Lake

(OTU 12).

E. Cabi 2248 Ankara: Sereflikochisar, 8 km to Sere-

flikochisar, shore of Tuz Lake (OTU 35).

E. Cabi 2247 Ankara: Sereflikochisar, 30 km to Sere-

flikochisar, shore of Tuz Lake (OTU 11).

E. bonaepartis

E. Cabi 339 Ankara: Polatlı to Sivrihisar, 26 km from

Polatlı (OTU20).

D28141 Sanlıurfa: S.Urfa to Akcakale, 13 km from

S.Urfa (OTU19).

E. Cabi 2241 Urfa: Ceylanpınar State Farm, Seyh Nasr

Streambed (OTU 17)

E. Cabi Urfa Ceylanpınar State Farm, edge of Gumussu

and Beyazkule, Sıhanlı Valley (OTU 18)

G. Akaydın 7764b Aksaray, Aksaray to Sereflikochisar,

around Tuz Lake 970 m (OTU 14)

E. Cabi 2255 Agrı: 9.5 km from Dogubeyazıt to Igdır(OTU 16)

E. Cabi 2240 Sanlıurfa: Ceylanpınar State Farm, E. of

Gellegec Karakolu (OTU 15)

E. distans

ANK 1459 Konya: Yavsan salt enterprise, nr Tuz Lake

(OTU 5).

E. Cabi 2256 Agrı: 9.5 km from Dogubeyazıt to Igdır(OTU 4).

E. Cabi 2506 Agrı: Agrı to Dogubeyazıt (OTU 3).

E. triticeum

E. Cabi 1706 Konya: Yavsan salt enterprise, near Tuz

Lake (OTU 1).

E. Cabi 2492 Erzurum: Horasan Town centre (OTU2).

References

Bentham G, Hooker JD (1883) Genera plantarum. Reeve, London

Bor NL (1968) Gramineae. In: Towsend CC, Guest E, El-Rawi A

(eds) Flora of Iraq, vol 9

Bor NL (1970) Gramineae. In: Rechinger KH (ed) Flora Iranica, vol

70. Graz, Austria: Akademische Druk-Und Verlagsanstalt, Wien

Cabi E, Dogan M (2009) A first vouchered wild record for the flora of

Turkey: Aegilops juvenalis (Thell.) Eig (Poaceae). Turk J Bot

33:447–452

Cabi E, Dogan M, Baser B, Us E, Pehlivan S (2009) Morphological

and palynological features of the Dasypyrum (Poaceae) in

Turkey. Phytol Balc 15:393–400

Cabi E, Dogan M, Mavi O, Karabacak E, Baser B (2010) Elymussosnowskyi (Hackel) Melderis (Poaceae), a rare endemic species

in Turkey. Turk J Bot 34:105–114

Clayton WD, Renvoize SA (1986) Genera graminum. Grasses of the

world. Kew Bull Addit Ser 13:155

Davis PH, Mill RR, Tan K (1988) Flora of Turkey and The East

Aegean islands. Vol. 10 (supplement). Edinburgh University

Press, Edinburgh

Dogan M (1988) A scanning electron microscope survey of lemma in

Phleum, Pseudophleum and Rhizocephalus (Gramineae). Notes

Roy Bot Gard Edinb 45(1):117–124

Dogan M (1991) Taxonomic significance of vegetative and floral

morphologies in the genus Alopecurus L. (Gramineae). Turk J

Bot 15:124–132

Dogan M (1992) Assessment of the morphological variaton by means

of numerical taxonomy in Alopecurus (Gramineae). Fl Veg

Mundi 9:75–81

Dogan M (1997) Numerical taxonomic study on the genus AlopecurusL. (Gramineae). Ot Sist Bot 4(2):71–76

Frederiksen S (1991) Taxonomic studies in Eremopyrum (Poaceae).

Nord J Bot 11:271–285

Frederiksen S (1993) Taxonomic studies in some annual genera of the

Triticeae (Poaceae). Nord J Bot 13:481–493

Frederiksen S (1994) Hybridization between Taeniatherum caputmedusae and Triticum aestivum. Nord J Bot 14:3–6

Frederiksen S, von Bothmer R (1989) Intergeneric hybridization

between Taeniatherum and different genera of Triticeae (Poa-

ceae). Nord J Bot 9:229–240

Gower JC (1971) A general coefficient of similarity and some of its

properties. Biometrics 27:857–874

Guner A, Ozhatay N, Ekim T, Baser KHC (2000) Flora of Turkey and

the East Aegean islands. Vol 11 (supplement). Edinburgh Univ

Press, Edinburgh

Holmgren PK, Holmgren NH, Barnett LC (1990) Index herbarium I.

The herbaria of the world. Regn Veg 120:1–693

Jaubert C, Spach E (1851) Ilustrationes plantarum orientalium, vol 4.

Paris

Kovach WL (1999) MVSP—a multivariate statistical package for

windows, version 3.1. Kovach Computing Services, Pentraeth

Ledebour CF (1853) Flora Rossica, vol 4. Stuttgart

Love A (1984) Conspectus of the Triticeae. Feddes Repert 95:425–521

Mason NWH, Mouillot D, Lee WG, Wilson JB (2005) Functional

richness, functional evenness and functional divergence: the

primary components of functional diversity. Oikos 111:112–118

Melderis A (1985) Eremopyrum (Ledeb.) Jaubert & Spach. In: Davis

PH (ed) Flora of Turkey and East Aegean Islands, vol 9.

Edinburgh, pp 227–230

Melderis A, Humpries CJ, Tutin TG, Heathcote SA (1980) Tribe

triticeae dumort. In: Tutin TG, Heywood VH, Burges NA,

Moore DM, Valentine DH, Walters SM, Webb DA (eds) Flora

Europaea, vol 5. Cambridge University Press, Cambridge,

England, pp 190–200


123

Nevski SA (1936) Conspectus Loliearum, Nardearum, Lepturearum

Hordeearumque Florae Unionis Rerum Publicarum Sovieticarum

Socialisticarum. Trudy Botani Eeskogo instituta Akademii nauk

SSSR. Ser. I. Flora i sistematika vyssih rastenij 2:33–90

Ozler H, Cabi E, Us E, Dogan M, Pehlivan S (2009) Pollen

morphology of Agropyron Gaertner in Turkey. Bangladesh J Pl

Taxon 16(1):21–28

Sakamoto S (1967) Genome analysis of the genus Eremopyrum.

Wheat Inform Serv 23–24:21–22

Sakamoto S (1968) Cytogenetic studies in the tribe Triticeae. VI.

Intergeneric hybrid between E. orientale and Aegilops squarrosa.

Jap J Genet 43:167–171

Sakamoto S (1972) Intergeneric hybridizations between E. orientale

and Henrardia persica, an example of polyploid species forma-

tion. Heredity 28:109–115

Sakamoto S (1974) Intergeneric hybridizations among three species

of Heteranthelium, Eremopyrum and Hordeum, and its signifi-

cance for the generic relationships within the Tribe Triticeae.

New Phytol 73:341–350

Sakamoto S (1979) Genetic relationships among four species of the

genus Eremopyrum in the tribe Triticeae, Gramineae. Mem Coll

Agric Kyoto Univ 114:1–27

Sneath PHA, Sokal RR (1973) Numerical taxonomy. W. H. Freeman,

San Francisco, California, USA

Tzelev NN (1976) Triticeae dumort. Poaceae URSS. Academia

Scientiarum URSS. Leningrad, pp 105–206


123

Taxonomic study on the genus Eremopyrum (Ledeb.) Jaub. et Spach (Poaceae) in Turkey

Documents

Transcript of Taxonomic study on the genus Eremopyrum (Ledeb.) Jaub. et Spach (Poaceae) in Turkey