PHYTOLOGIA BALCANICA 15 (3): 393 – 400 Sofia, 2009 393 Morphological and palynological features of the genus Dasypyrum (Poaceae) in Turkey Evren Cabi 1*, Musa Doğan1, Birol Başer2, Ebru Us2 & Sevil Pehlivan2 1 Department of Biological Sciences, Faculty of Arts and Sciences, Middle East Technical University, Ankara, Turkey, e-mail: ecabi@metu.edu.tr, ecabi2004@yahoo.com (author for correspondence*) 2 Department of Biology, Gazi University, Faculty of Arts and Science, Teknikokullar, 06500 Ankara, Turkey Received: June 22, 2009 ▷ Accepted: October 13, 2009 Abstract. This article presents a revision of the genus Dasypyrum in Turkey, based on 25 populations gathered during field trips between 2006 and 2008. These populations were subjected to examination on the basis of their morphological and palynological characteristics. A corrected and updated description, as well as a distribution map of D. villosum are provided. Furthermore, the relationship between Dasypyrum and two other genera included in the tribe Triticeae (Secale and Triticum) is discussed on the basis of palynological features. The palynological investigations showed that the genus Dasypyrum is close to Secale rather than to Triticum. Key words: Dasypyrum, distribution, pollen morphology, Triticeae, Turkey Introduction Genus Dasypyrum (Coss. & Durieu) T. Durand (Poaceae) is considered a very important genetic source in breeding experiments, due to its close phylogenetic relationship with Triticum L. and Secale L., and particularly their cultivated species. The genus is represented with two species: D. villosum (L.) P. Candargy and D. breviaristatum (H. Lindb.) Fred. The former is an annual diploid species distributed naturally in Turkey. The latter perennial species consists of diploid and tetraploid cytotypes and is distributed only in a restricted area over 1000 m a.s.l. in Morocco and Greece. Morphologically, this genus is quite distinct from the other genera within the tribe Triticeae by its two-keeled glumes (Frederiksen 1991; Galasso & al. 1997; Ohta & Morishita 2001). Dasypyrum villosum was originally named Secale villosum by Linnaeus (1753). Later on it was referred 13 • Phytol. Balcan. 15(3) • 2009 to various genera, such as Agropyron Gaertn., Haynaldia Schur and Triticum. On the basis of morphological similarities, many researchers refer to Dasypyrum is closer to the Triticum/ Agropyron complex and Secale (Sakamoto 1973; West & al. 1988). According to Kellogg (1989), Dasypyrum was close to Agropyron Gaertn., Crithodium Link. and Eremopyrum (Ledeb.) Jaub. & Spach, because they all have keeled glumes. On the other hand, Kellogg (1992) showed that V genome of D. villosum was closely related to S genome of Pseudoroegneria (Nevski) Á. Löve and Elymus repens (L.) Gould. on the basis of cpDNA. Also, on the basis of the chloroplast DNA data, Mason Gamer (2004) showed that D. villosum is closely associated with Pseudoroegneria, Elymus L. and Thinopyrum Á. Löve. This phylogenetic view was equally supported by Hsiao & al. (1995) on the basis of the sequences of an internal transcribed spacer (ITS) region of the nuclear ribosomal DNAs of D. villosum and Pseudoroegneria. 394 Cabi, E. & al. • Morphology and palynology of Dasypyrum in Turkey The relationship between the genera Dasypyrum and Triticum has been investigated in various ways, such as polymorphism of storage proteins (Montebove & al. 1987; Shewry & al. 1987), isozymes points (Montebove & al. 1987; Liu & al. 1995) and crossing efforts (Blanco & al. 1988; Bothmer & Claesson 1990). The above-mentioned studies suggested a closer relationship with Triticum. On the other hand, morphological (Baum 1977, 1978a, b), biochemical (Baum 1983), cytological, and molecular DNA analyses (Linde-Laursen & al. 1992; Vershinin & HeslopHarrison 1998) and DNA/DNA hybridization experiments (Lucas & Jahier 1988; Uslu & al. 1999) suggested a closer relationship with Secale. Several phylogenetic analyses of the genus Dasypyrum have been reported so far on the basis of morphology, cytology and molecular biology (Gradzielewska 2006). Relatively few studies have been conducted on the genus Dasypyrum and its morphologically close genera Triticum and Secale regarding their pollen morphology (Fritsch & al. 1977; Kruse 1980). In this paper, in an attempt to revise the genus Dasypyrum in Turkey and to contribute to the phylogeny of this genus in the tribe, palynological features of D. villosum and its morphologically close species T. monococum L. and S. cereale L. are compared. Material and methods The specimens used in both morphological and palynological parts of our study are collected from natural habitats of the species in Turkey. Voucher specimens are kept in the GAZI herbarium. The description of D. villosum is based on measurements of about 130 specimens representing 25 D. villosum populations collected from Turkey (Table 1). Morphological measurements were made with Leica L2 Stereomicroscope and Leica Application Suite software package. Sterile parts of the specimens were measured directly on pressed material and the results were given in a range in the description; fertile parts are measured after boiling in tap water. The measurements of fertile parts of D. villosum include: spike length (SpL), spikelet length (SpkL), glume length (GL), lemma length (LL), palea length (PL) awn length of glumes and awn length of lemmas. Five randomly taken spikes from each population were measured. Means and standard deviations (SD) were computed for each population by means of Microsoft Excel (Table 2). Table 1. List of studied populations of D. villosum. / No Collector Number 1 E. Cabi 3183 2 E. Cabi 3173 3 E. Cabi 3199a 4 5 6 7 8 9 10 11 E. Cabi 783 E.Cabi 747 E.Cabi 752 E.Cabi 763 E. Cabi 801 E. Cabi 848 E. Cabi 774 E. Cabi 3162 12 E .Cabi 722 13 E Cabi 857 E. Cabi 817 E. Cabi 3204 E. Cabi 3210 E. Cabi 805 E. Cabi 515 E. Cabi 525 E. Cabi 304 E. Cabi 516 E. Cabi 521 E. Cabi 533 E. Cabi 537 14 15 16 17 18 19 20 21 22 23 24 25 Northing/ Easting A1: Edirne: Keşan to Enez, 30 km to 40°40.571'N Enez, 132 m, 03 June 2008. 26°20.785'E A1: Çanakkale: Gelibolu to Keşan, 8 km 40°45.978'N to Keşan, roadsides, 50 m, 03 June 2008. 26°42.213'E A1 Edirne: Enez to İpsala, 2 km N. of 40°44.033'N Enez towards the border with Greece, 26°07.435'E 31 m, 03 June 2008. A1: Kırklareli: Hasköy to Kırklareli, 41°40.829'N 167 m, roadsides, 29 May 2006. 27°06.208'E A1: Tekirdağ: Gaziköy to Uçmakdere 40°47.402'N arası, 127 m, 28 May 2006. 27°21.997'E A1 Tekirdağ: Uçmakdere to Kumbağ, 40°48.788'N 276 m, roadsides, 28 May 2006. 27°23.387'E A1 Tekirdağ: Gaziköy to Kumbağ, 5 km 40°50.472'N to Kumbağ, 234 m, open sides of Quercus, 27°26.101'E Pinus, 28 May 2006. A1 Kırklareli:Vize to Saray, Çakıllı köyü 41°28.571'N yakınları, 184 m, roadsides, 29 May 2006. 27°53.249'E A1 Tekirdağ: Şarköy to Gelibolu, 91 m, 40°34.301'N 30 May 2006. 26°56.103'E A1 Tekirdağ: Tekirdağ to Silivri, 20 km 41°00.787'N from Tekirdağ, 7 m, 27°44.278'E 40°12'56"N A1(E) Çanakkale: Eceabat, 36 km to 26°22'81"E Gelibolu, next to beach under Kavak ağacı, 0-5 m, 03 June 2008. A1(E) Çanakkale Eceabat-Gelibolu pe- 40°15.210'N ninsula, near Anzac Newzealand outpast, 26°16.819'E 8 m, maritime sands, 28 May 2006. A1(A) Çanakkale: İntepe, 127 m., road- 40°00.534'N sides, 31 May 2006. 26°19.534'E A2(E) İstanbul: Silivri to Tekirdağ, 41°00.171'N around German sitesi, 23 m, dry pastures. 27°58.388'E A1(E) İstanbul: Silivri, s.l., 04 June 2008. 41°04.128'N 28°08.468'E A2 (E) Belgrad forest, 05 June 2008. Location A1 (E) Halkalı train station environs, 21 m, 30 May 2006. B1 İzmir: Tire to Selçuk, 1.5 km to Belevine, 12 May 2006. C1 Aydın Selçuk to Şirince, 2 km to Şirince, 250 m, under olive trees, 12 May 2006. C1 Muğla: Milas to Akbüke, before 7 km to Kızılağaç village, 160 m, 04 May 2006 C1 Aydın Selçuk to Efes, 7 m, roadsides, 12 May 2006. C1 Aydın Efes next to historical sites of Efes, 8 m, 12 May 2006. C1 Aydın: Selçuk to Kuşadası, 89 m, 13 May 2006. C1 Aydın: Kuşadası Güzel Çamlı, National Park of Dilek peninsula, 65 m, 13 May 2006. E. Cabi C2 Denizli: NW of Denizli, Üçler, 31 3824 May 2008. 41°01.112'N 28°046.081'E 38°01.004'N 27°28.304'E 37°19.418'N 27°36.410'E 37°56.856'N 27°20.541'E 37°56.690'N 27°20.514'E 37°55.034'N 27°16.788'E 37°42.401'N 27°12.119'E 37°47.706'N 29°01.064'E Phytol. Balcan. 15(3) • Sofia • 2009 395 Table 2. Morphological measurements of D. villosum accessions gathered during field studies (mean + standard deviation) (n = 5). No Collector / Number SpL (cm) SpkL (mm) GL (mm) LL (mm) PL (mm) AofG (mm) AofL (mm) 1. E .Cabi 3183 6.2±0.57 14.2±1.09 5.5±0.50 10.9±0.74 11.2±0.83 3.74±0.25 4.04±0.36 2. E. Cabi 3173 7.2±0.57 20.2±0.83 6.2±0.44 11.6±1.14 11.3±0.83 3.06±0.56 3.68±0.21 3. E. Cabi 3199a 5.0±0.35 20.4±1.14 7.2±0.27 12.0±0.70 11.4±0.41 4.54±0.36 5.8±0.27 4. E. Cabi 783 4.7±0.25 16.5±1.22 6.0±0.21 13.0±0.70 12.1±0.41 4.86±0.21 5.4±0.41 5. E. Cabi 747 5.2±075 15.4±0.54 5.9±0.22 11.3±0.44 11.0±0.35 4.5±0.35 5.1±0.74 6. E. Cabi 752 6.9±0.74 15.0±0.70 6.4±0.54 11.0±0.70 10.5±0.50 6.02±0.31 6.6±0.41 7. E. Cabi 763 6.4±0.41 16.0±0.70 6.2±0.57 10.6±0.54 10.4±0.41 4.4±0.65 5.1±0.54 8. E. Cabi 801 7.4±0.89 14.0±0.70 6.0±0.36 12.2±0.44 12.0±0.70 4.06±0.56 4.5±0.35 9. E. Cabi 848 6.2±0.57 17.1±0.54 5.9±0.54 11.8±0.83 11.1±0.54 4.1±0.41 4.5±0.25 10. E. Cabi 774 7.3±1.20 18.6±1.14 6.5±0.35 14.0±0.70 13.3±0.44 3.6±0.41 5.2±0.57 11. E. Cabi 3162 5.5±1.11 15.6±1.14 5.9±0.21 11.5±0.5 11.3±0.57 4.2±0.23 4.42±0.42 12. E. Cabi 722 5.5±0.50 15.2±1.30 5.8±0.27 11.3±0.44 11.2±0.44 5.0±0.79 4.4±0.41 13. E. Cabi 857 5.7±1.20 19.2±0.83 5.7±0.44 12.8±0.83 12.0±0.70 4.1±0.56 4.50±0.35 14. E. Cabi 817 6.1±0.89 19.7±0.44 6.7±0.44 12.9±0.5 12.5±0.5 6.1±0.41 5.9±0.54 15. E. Cabi 3204 5.7±0.65 20.4±1.14 6.5±0.35 12.0±0.70 11.1±0.54 5.5±0.5 5.1±0.54 16. E. Cabi 3210 3.5±0.39 18.4±0.89 6.2±0.57 11.0±0.70 10.5±0.50 5.2±0.57 5.4±0.41 17. E. Cabi 805 7.4±0.89 18.4±0.89 5.8±0.27 12.5±0.5 12.0±0.70 5.5±0.5 5.9±0.54 18. E. Cabi 515 5.5±1.65 17.1±0.54 5.5±0.35 11.5±0.5 11.2±0.44 4.4±1.08 5.2±0.57 19. E. Cabi 525 4.7±0.97 20.4±1.14 5.5±0.35 12.0±0.70 11.2±0.44 3.4±1.14 5.4±0.41 20. E.Cabi 304 4.0±0.70 16.3±0.83 5.9±0.21 9.6±0.65 9.5±0.5 4.7±1.64 5.9±0.54 21. E. Cabi 516 8.2±0.54 19.3±0.44 6.5±0.35 13.0±0.70 12.7±0.44 3.7±0.25 3.7±0.25 22. E. Cabi 521 3.8±0.23 17.1±0.54 6.7±0.44 12.0±0.70 11.1±0.54 3.1±0.56 3.9±1.24 23. E. Cabi 533 4.5±0.35 18.4±0.89 6.7±0.44 11.5±0.5 11.2±0.44 4.1±0.41 4.7±1.64 24. E. Cabi 537 6.1±0.89 20.2±0.83 5.9±0.54 13.5±0.5 12.7±0.44 4.7±1.64 5.2±0.57 25. E. Cabi 3824 5.5±0.50 17.1±0.54 6.2±0.57 13.5±0.5 12.5±0.50 3.4±1.14 4.5±0.25 SpL: spike length; SpkL: spikelet length; GL: glume length; LL: lemma length (mm); PL: palea length (mm); AofG: awn length of glume (mm); AofL: awn length of lemma. Pollen samples of each studied taxon were Table 3. List of palynologicaly studied accessions. obtained from the herbarium species listed in Collector / Northing / No Number Taxa Locality Easting Table 3. For palynological investigations, pollen 1. E.Cabi Dasypyrum C1 Aydın: Selçuk to Efes, 7 m, 37°56.856'N slides for morphological examination by light 516 villosum roadsides, 12 May 2006. 27°20.541'E microscopy were prepared according to Wo2. E. Cabi Secale B2 Uşak: Karahallı to Ulubey, 1 38°24.304'N 2244 cereale var. km to Ulubey, edge of cultivated 29°17.394'E dehouse (1935) and Erdtman (1952), and the cereale fields; 712m. 11 May 2006. measurements were made with a Leica DM1000 3. E. Cabi S. cereale var. B10 Ağrı: Doğubeyazıt to Iğdır, 9 39°38.554'N microscope and Leica Application Suite soft2545 vavilovii km to Iğdır, roadsides; 1589 m, 08 44°02.178'E July 2007. ware package. Measurements include the fol4. E. Cabi Triticum A4 Kastamonu: İhsangazi, Kuş- 41°14'29.8"N lowing parameters: long axis of spheroidal pol2374 monococcum çalar district, Sekicek street, lime- 33°31'39.1"E stone slopes, 900 m, 13 June 2007. len grains (A), short axis of spheroidal pollen grains (B), long axis of elipsoid porus (pa), short axis of elipsoid porus (pb), exine thickness, the Department of Biology, Gazi University. SEM annulus diameter (An), A/B ratio, operculum diimages were used to describe surface texture of the ameter (Op.), thickest part of intine (I), and intine pollen. Scabrae density per 1 μm2, as well as the thickness (i) (Table 4). height and width of scabrae are measured directIn order to study the pollen texture ornamentaly on the SEM images, using the facilities of imtion, dry pollen grains were first mounted on douage processing software Adobe Photoshop CS2. ble-sided carbon tape affixed to aluminum stubs. Ten randomly chosen 1 μm2 areas and scabrae on Grains were coated with gold with a Hummle VII the image were measured. The descriptive termisputter coater and observations were made using a nologies of Faegri & Iversen (1975) and Punt & al. Jeol 6060 Scanning Electron Microscope (SEM) at (2007) were followed. 396 Cabi, E. & al. • Morphology and palynology of Dasypyrum in Turkey Table 4. Morphology of pollen parameters (μm) of the three investigated taxa (mean + standard deviation) (n = 30). Taxon Daspyrum villosum Preparation A B A/B Pa Pb I i Exine Op An W 39.39±1.86 33.67±2.17 1.19 3.76±0.64 3.22±0.36 3.74±0.51 0.56±0.12 1.34±0.31 2.60±0.61 8.36±0.27 E 52.19±3.76 40.32±2.76 1.29 4.45±0.50 3.99±0.68 1.80±0.30 10.05±0.36 Secale cereale. W 42.30±4.07 33.72±3.46 1.25 4.25±0.38 3.62±0.27 3.62±1.06 0.87±0.33 1.24±0.21 1.77±0.19 7.70±0.57 var. cereale E 40.17±1.86 30.03±1.90 1.33 3.48±0.26 3.15±0.29 1.34±0.17 8.16±0.58 Triticum W 35.34±1.83 35.34±1.17 1.00 3.69±0.33 3.45±0.34 3.41±1.21 0.82±0.12 1.59±0.16 1.99±0.27 8.89±0.80 monococcum E 32.72±1.99 32.38±2.03 1.01 3.27±0.20 2.92±0.21 1.55±0.18 8.84±0.54 W: non-acetolyzed pollen grains; E: acetolysed pollen grains; A: long axis of spheroidal pollen grains; B: short axis of spheroidal pollen grains; A/B: shape of pollen grains; Pa: long axis of ellipsoid porus; Pb: short axis of ellipsoid porus; I: thickest part of the intine; i: intine thickness; Op: operculum diameter; An: annulus thickness. Results and discussion Dasypyrum villosum Plant morphology (Plate I, Figs 1–6) Tufted annual. Stems 25–100 cm with 2–5 nodes, usually decumbent or geniculate-ascending, glabrous. Leaf blades 7–22 cm × 2–4 mm, flat, sparsely scabrous to densely pilose on the upper surface, glabrous beneath. Spike 4–10 cm; rachis densely barbate on margins. Spikelets 7–20 mm (excl. awn), compressed. Glumes 5–7.5 mm long, with small fascicles of rigid hairs 2–4 mm long on keels, with 2.0–5 cm scabrid awn. Lower lemma 10–16 mm, glabrous below, sparsely pilose above, with fascicles of rigid hairs below awn; awn 2–6 cm, scabrid. Palea 10–15 mm. long, membranous, anthers 5–8 mm long, caryopsis 4–5 mm long. Habitat and ecology. The species grows in sandy arid areas, on rocky limestone slopes. Its altitudinal range varies from sea level to 200 m. Other species growing in Fig. 1. Distribution of D. villosum in Turkey. the area with D. villosum include: Cynosurus cristatus L., Secale cereale L., Taeniatherum caput-medusae (L.) Nevski., Hordeum murinum Huds., Hordeum bulbosum L., H. murinum L., Elymus elongatus (Host) Runemark, Quercus coccifera L., Cistus creticus L., Stachys cretica subsp. lesbiaca Rech. f., Bromus sterilis L., B. tectorum L., Aegilops geniculata Roth, A. triuncialis L., Psoralea bituminosa L., Trifolium campestre Schreb., T. arvense L., Ononis viscosa L., Onobrychis caput-galli Lam., Thymbra spicata L., Coridothymus capitatus (L.) Rchb. f., Sonchus oleraceus L., Juniperus oxycedrus L., Salvia viridis L., S. virgata Jacq., Hymenocarpus circinnatus Savi, Sarcopoterium spinosum Spach, Avena fatua L. Phenology. Flowering in May to June, fruiting from June to July. General distribution. European Turkey, W. Anatolia, S. European islands, S.W. Russia, Crimea, Caucasia. A Mediterranean element. The species is confined from A1 (A1 sensu Davis 1965) to C2 squares in Turkey (Fig. 1). Phytol. Balcan. 15(3) • Sofia • 2009 Palynological features (Plates II & III) The results of palynological investigations have shown that all pollen grains of D. villosum and its morphologically similar species S. cereale (wild and cultivated varieties) and T. monococum are monoporate and have scabrate exine surfaces. The pollen shape and scabra density per 1 μm2, as well as the height and width of scabrae are specific features for differentiation of the taxa. The mean and standard 397 deviations of the measured pollen parameters of taxa are given in Table 4. All investigated taxa had monoporate and prolate, subprolate or prolate spheroidal pollen grains. The pore was surrounded by an annulus and was partly covered by an operculum. While in D. villosum and S. cereale the pollen shape is subprolate (1.14–1.33 μm) or prolate (1.33–2.00 μm), in T. monococum the pollens are prolate spheroidal (1.00–1.14 μm) (Plate II, Figs 1–4; Table 4). Plate I Figs 1–6. Morphological features of D. villosum: 1, Spike; 2, Close view of spike; 3, Glume; 4, Indumentum of leaf blades, 5, Seed; 6, Palea and seed. 398 Cabi, E. & al. • Morphology and palynology of Dasypyrum in Turkey Pollen grains of Poaceae were classified as annulate or nonannulate, and operculate or nonoperculate by Salgado-Labouriau & al. (1990), Chaturvedi & al. (1998) and Perveen (2000, 2006). In this study, we have observed that all examined taxa were annulate and operculate. The exine ornamentation types in this family were defined as insular, granulose, spinulose, verrucose, and brevicerebro ornate (Chaturvedi & al. 1994, 1998; Liu & al. 2004). Erdtman (1969), Moore & al. (1978, 1991), Başer & al. (2009), and Özler & al. (2009) used the term scabrate for exine sculpture covered with small (<1 μm) elements, equivalent to granulate exine ornamentations. According to our SEM investigations, exine sculpture is scabrate in all examined taxa (Plate III, Figs 1–3). Kohler & Lange (1979) suggested that the number of spinules may be used for identification. In the present study, we have found that the investigated taxa could be differentiated on the basis of the exine parameters, such as the number and distance between the scabrae. In S. cereale var. cereale, the number of scabrae in 1 μm2 varies from 5 to 10 (8.8±2.18); the width of scabrae is 0.40±0.12 μm and the height of scabrae is 0.50 ± 0.16 μm. In T. monococum, the number of scabrae in 1 μm2 is 12–15 (13.5±1.15); the width of scabrae is 0.30±0.08 μm and the height of scabrae is 0.30±0.09 μm. In D. villosum, the number of scabrae in 1 μm2 is 3–5 (4.5±0.5); the width of scabrae is 0.30±0.08 μm and the height of scabrae is 0.1±0.06 μm (Plate III, Figs 1–3). The potential usefulness of the pollen exine features of the Gramineae as markers of gene expression and inheritance was shown mainly in intergeneric hybrids, where the influence of one or both parents on the exine sculpture of the resulted hybrids was recorded (Grant 1972; Rajendra & al. 1978). The pollen shape and density of scabrae no/μm2 on the exine surface can be used as the most functional differentiating characters. On the basis of pollen shape and the number of scabrae Dasypyrum is closer to Secale rather than to Triticum. Acknowledgements. The authors wish to extend their thanks to the Scientific and Technical Research Council of Turkey (TUBITAK-TBAG-105 T 171) for its financial support. The authors are also grateful to the anonymous reviewers for the critical notes and suggestions. Plate II Figs 1–4. Light micrographs of pollen grains of: 1, D. villosum; 2, S. cereale; 3. S. cereale var. vavilovi; 4, T. monococum. 399 Phytol. Balcan. 15(3) • Sofia • 2009 Plate III Figs 1–3. SEM observations of: 1, D. villosum, a – general view; b – exine ornamentation; 2, S. cereale, a – general view; b – exine ornamentation; 3, T. monococum, a – general view; b – exine ornamentation. References Başer, B., Özler, H., Cabi, E., Doğan, M. & Pehlivan, S. 2009. Pollen morphology of the genus Eremopyrum (Poaceae) in Turkey. – World Appl. Sci. J., 6(12): 1655-1659. Baum, B.R. 1977. Taxonomy of the tribe Triticeae (Poaceae) using various numerical techniques I. – Can. J. Bot., 55: 1712. Baum, B.R. 1978a. Taxonomy of the tribe Triticeae (Poaceae) using various numerical techniques. II. – Can. J. Bot., 56: 27. Baum, B.R. 1978b. Genetic relationships in Triticeae based on computations of Jardine & Sibson Bk clusters. – Can. J. Bot., 56: 2948. Baum, B.R. 1983, Phylogenetic analysis of the tribe Triticeae (Poaceae) based on the morphological characters of the genera. – Can. J. Bot., 61: 518-535. Blanco, A., Perrone, V. & Simeone, R. 1988. Chromosome pairing variation in Triticum turgidum L., Dasypyrum villosus (L.) Candargy hybrids and genome affinities. – In: Miller, T.E. & 400 Cabi, E. & al. • Morphology and palynology of Dasypyrum in Turkey Koebner, R.M.D. (eds), Proc of the 7th Int. Wheat Genetic Symp., Cambridge, England, July 13-19. Vol. 1, pp. 63-67. 8th Int. Wheat Genetic Symp. Pp. 181-185. Chinese Agricultural Sci. Press, Bejing. Bothmer, von R. & Claesson, L. 1990. Production and meiotic pairing of intergeneric hybrids of Triticum and Dasypyrum species. – Euphytica, 51: 109-117. Liu, Q., N.-X, Zhao, & Hao, G. 2004. Pollen morphology of the Chloriodoideae (Gramineae). – Grana, 43: 238-248. Chaturvedi, M., Yunus D. & Datta, K. 1994. Pollen morphology of Sorghum Moench sections Eu- sorghum and Para-sorghum. – Grana, 33: 117-123. Chaturvedi, M., Datta, K. & Nair, P.K.K. 1998. Pollen morphology of Oryza, Poaceae. – Grana, 37: 79-86. Davis, P.H. (eds). 1965. Flora of Turkey and the East Aegean Islands. Vol. 1. Edinburgh Univ. Press, Edinburgh. Erdtman, G. 1952. Pollen Morphology and Plant Taxonomy. Angiosperms. (An introduction to Palynology I). Almqvist & Wiksell, Stockholm. Erdtman, G. (ed.) 1969. Handbook of Palynology, Hafner Publishing Co., New York. Faegri, K. & Iversen, J., 1975. Textbook of Pollen Analysis. Hafner Press, New York. Frederiksen, S. 1991. Taxonomic studies in Dasypyrum (Poaceae). – Nord. J. Bot., 11: 135-142. Fritsch, R., Kruse, J., Ohle, H. & Schaefer, H. 1977. Anatomical studies within the genera Triticum L. and Aegilops L. (Gramineae). – Kulturpflanze, 25: 155-265. Galasso, I., Blanco, A., Katsiotis, A., Pignone, D. & HeslopHarrison, J.S. 1997. Genome organization and phylogenetic relationships in the genus Dasypyrum analysed by Southern and in-situ hybridization of total genomic and cloned DNA probes. – Chromosoma, 106: 53-61. Gradzielewska, A. 2006. The genus Dasypyrum – part 1. The taxonomy and relationships within Dasypyrum and with Triticeae species. – Euphytica, 152: 429-440. Grant, C.A. 1972. A scanning electron microscopy survey of some Maydeae pollen. – Grana, 12: 177-184. Hsiao, C, Chatterton, N.J., Asay, K.H. & Jensen, K.B. 1995. Phylogenetic relationships of the monogenomic species of the tribe Triticeae (Poaceae) inferred from nuclear rDNA (internal transcribed spacer) sequences. – Genome, 38(2):211-223. Kellogg, E.A. 1989. Comments on genomic genera in the Triticeae (Poaceae). – Am. J. Bot., 76: 796-805. Kellogg, E.A. 1992 Tools for studying the chloroplast genome in the Triticeae (Gramineae): An EcoRI map, a diagnostic deletion, and support for Bromus as an outgroup. – Am. J. Bot., 7(2):186197. Kruse, J. 1980. Exine ornamentations of the genera Triticum L. and Aegilops L. – Kullturpflanze, 28: 341-359. Linde-Laursen, I., von Bothmer, R. & Jacobsen, N. 1992. Relationships in the genus Hordeum: Giemsa C-banded karyotypes. – Hereditas, 116: 111-116. Linnaeus, C. 1753. Species Plantarum, ed. 1. Stockholm. Liu, D.J., Chen, P.D. & Raupp, W.J. 1995. Determination of homoeologous groups of Haynaldia villosa chromosomes. – In: Proc. Lucas, H. & Jahier, J. 1988. Phylogenetic relationships in some diploid species of Triticineae: cytogenetic analysis of interspecific hybrids. – Theor. Appl. Genet., 75: 498-502. Mason-Gamer, RJ. 2004. Reticulate evolution, introgression, and intertribal gene capture in an allohexaploid grass. – Syst. Biol., 53(1): 25-37. Montebove, L., De Pace, C., Jan, C.C., Scarascia-Mugnozza, G.T. & Qualset, C.O. 1987. Chromosomal location of isozyme and seed storage protein genes in Dasypyrum villosum (L.) Candargy. – Theor. Appl. Genet., 73:836-845. Moore, P.D. & Webb, J.A. (eds). 1978. An Illustrated Guide to Pollen Analysis. Hodder and Stoughton, London. Moore, P.D., Webb, J.A. & Collinson, M.E. 1991. Pollen Analysis. 2nd Ed. Blackwell Sci. Publications, Osney Mead, Oxford. Ohta, S. & Morishita, M. 2001. Genome relationships in the genus Dasypyrum (Gramineae). – Hereditas, 135: 101-110. Özler, 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. Perveen, A. 2000. Pollen characters and their evolutionary significance with special reference to the Flora of Karachi. – Türk. Biyol. Derg., 24: 365-378. Perveen, A. 2006. A Contribution to the pollen morphology of family Gramineae. – World Appl. Sci. J., 2: 60-65. Punt, W., Hoen, P.P., Blackmore, S., Nilsson, S. & Le Thomas, A. 2007. Glossary of pollen and spore terminology. – Rev. Palaeobot. Palynol., 143(1-2): 1-81. Rajendra, B.R., Tomb, A.S., Mujeeb, K.A. & Bates, L.S. 1978. Pollen morphology of selected Triticeae and two intergeneric hybrids. – Pollen & Spores, 20(1): 145-156. Sakamoto, S. 1973. Patterns of phylogenetic differentiation in the tribe Triticeae. – Rep. Kihara Inst. Biol. Res., 24: 11-31. Salgado-Labouriau, M.L & Rinaldi, M. 1990. Palynology of Gramineae of the Venezuelan Mountains. – Grana, 29: 119-128. Shewry, P.R., Parmar, S. & Pappin, D.J. C. 1987. Characterization and genetic control of the prolamins of Haynaldia villosa: relationship with cultivated species of the Triticeae (rye, wheat and barley). – Biochem. Genet., 25:309-325. Uslu, E, Reader, S.M. & Miller, T.E. 1999. Characterization of Dasypyrum villosum (L.) Candargy chromosomes by fluorescent in situ hybridization. – Hereditas, 131:129-134. Vershinin, A.V. & Heslop-Harrison, J.S. 1998. Comparative analysis of the nucleosomal structure of rye, wheat and their relatives. – Pl. Molec. Biol., 36:149-161. West, J.G., McIntyre, C.L. & Appels, R. 1988. Evolution and systematic relationships in the Triticeae (Poaceae). – Pl. Syst. Evol., 160: 1-28. Woodehouse, R.P. 1935. Pollen Grains. Hafner, New York.