Agro-Morphological Characterization of Some Wild Wheat (Aegilops L. and Triticum L.) Species

Turk J Agric For30 (2006) 387-398© TÜB‹TAK

387

Agro-Morphological Characterization of Some Wild Wheat(Aegilops L. and Triticum L.) Species

Alptekin KARAGÖZ, Necmi P‹LANALI, Turgay POLAT

Central Research Institute for Field Crops, P.O. Box: 226, 06042 Ankara - TURKEY

Received: 20.03.2006

Abstract: Wild wheat (Aegilops L. and Triticum L.) populations were collected from different regions of Turkey and characterizedfor 5 basic traits, in order to study their agro-morphological characteristics and variation among populations. Material of the studyconsisted of 112 populations of wild wheat and 12 populations of cultivated wheat. Cultivated einkorn (T. monococcum L. subsp.monococcum) and cultivated emmer (T. turgidum L. subsp. dicoccon (Shrank) Thell.) were used as checks for Triticum species.Among the Aegilops material studied, Ae. tauschii Coss. var. meyeri (Griseb. ex Ledeb.) has not been defined in the flora of Turkeybefore and is characterized agromorphologically for the first time in this study. Materials were transplanted to Haymana-AnkaraResearch Farm of the Central Research Institute for Field Crops after being germinated in a greenhouse in fall 2002. Populationswere characterized for plant height, number of days to heading, growth habit, number of stems per plant and spike length. Allobservations and measurements were performed on the same randomly selected plants. Mean values and coefficient of variationwere computed for all the populations. The highest variation was recorded for number of stems, while the lowest was for planthabit. Clusters based on a Euclidian similarity matrix ranked the species according to the sections they are located in. Considerablevariation was observed, even between populations collected from nearby sites. It was concluded that existing ex-situ conservationprograms should take into consideration that fact by sampling over short distances in order to capture as much of the variation aspossible.

Key Words: Wild wheat, Aegilops, Triticum, characterization, variation

Kimi Yabani Bu¤day (Aegilops L. ve Triticum L.) TürlerininAgro-Morfolojik Karakterizasyonu

Özet: Yabani bu¤day (Aegilops L. ve Triticum L) populasyonlar› Türkiye’nin de¤iflik bölgelerinden toplanarak, befl temel özelli¤i vepopulasyonlar aras› farkl›l›klar› ortaya koymak amac›yla agro-morfolojik yönden karakterize edilmifltir. Bu¤day›n 112 yabani ve 12kültür örne¤i bu çal›flman›n materyalini oluflturmufltur. Yabani siyez (T. monococcum L. subsp. monococcum) ve yabani gernik(T. turgidum L. subsp. dicoccon (Shrank) Thell.) örnekleri, Triticum türleri için kontrol olarak kullan›lm›flt›r. Materyal içinde yer alanAe. tauschii Coss. var. meyeri (Griseb. ex Ledeb.) türü Türkiye’nin floras›nda daha önce tan›mlanmam›fl olup ilk kez bu çal›flmaylaagromorfolojik karakterizasyonu yap›lm›flt›r. Tohum örnekleri 2002 y›l› sonbahar›nda serada çimlendirildikten sonra, Tarla BitkileriMerkez Araflt›rma Enstitüsü’nün Haymana’daki Araflt›rma ve Üretme Çiftli¤ine dikilmifltir. Populasyonlar bitki boyu, baflaklanma günsay›s›, geliflme formu, bitki bafl›na sap say›s› ve baflak uzunluklar› bak›m›ndan karakterize edilmifltir. Tüm ölçüm ve gözlemler,rastgele seçilen 5 bitki üzerinde yürütülmüfltür. Her populasyon için ortalamalar ve varyasyon katsay›lar› hesaplanm›flt›r. En yüksekde¤iflim katsay›s› sap say›s›nda, en düflü¤ü ise geliflme formunda bulunmufltur. Kümeleme Analizi türlerin kendi özel gruplar›nda yerald›¤›n› göstermifl, birbirine çok yak›n alanlardan toplanan örnekler aras›nda bile farkl›l›klar oldu¤u saptanm›flt›r. Bu durumda ex-situkoruma programlar›n›n mümkün olan en genifl varyasyonu yakalayabilmeleri için k›sa mesafelerde örnekleme yapmalar›n›n gereklioldu¤u sonucuna var›lm›flt›r.

Anahtar Sözcükler: Yabani bu¤day, Aegilops, Triticum, karakterizasyon, de¤iflim

* Correspondence to: [email protected]

Introduction

Rapid changes in climatic and environmentalconditions constitute significant threats to cultivatedcrops for adaptation to newly emerging conditions. Dueto universal problems such as air and soil pollution,depletion of the ozone layer, and global warming, it isinevitable that popular cultivars of today are not likely tobe able to cope with these challenges in the near future.Stress conditions played a major role in the evolution ofliving things. Wild relatives of cultivated crops that havebeen sustaining under intensive stress conditions forthousand of years, by modifying themselves to adapt tonewly emerging conditions, constitute a considerablesource (Nevo et al., 2002). Present conditions call forurgent measures. The study of plant material withdesired traits through characterization programs is anessential step for effective utilization of crop germplasm.

Long before the development of sophisticatedmethods for gene introgression between wild andcultivated, Gökgöl (1935) reported that northernEuropean countries had already begun to test the wildrelatives in their breeding programs.

Playing a significant role in feeding the worldpopulation, wheat is considered a fundamental crop.Wheat is assumed to have originated from the FertileCrescent (Harlan, 1981). Wild relatives of wheat are quitewidespread in Turkey, especially in the southeast. It isagreed that diploid wheat was first cultivated inKaracada¤, which is located in southeast Turkey, and soondispersed to the other parts of the world (Heun et al.,1997; Diamond, 1997; Nesbit and Samuel, 1998; Lev-Yadun et al., 2000). Triticum L. and Aegilops L. haveseveral species in 3 ploidy levels, namely diploid (2n = 14),tetraploid (2n = 28) and hexaploid (2n = 42). Davis(1985) reported that 16 Aegilops and 4 wild Triticumspecies exist in Turkey. Kimber and Feldman (1987) statethat the number of wild wheat relatives is 27 in the world,20 of which are in Turkey. Having adapted to a variety ofharsh conditions accompanied by heavy grazing bydomestic animals, wild wheat species have sustained theirdiversity to date.

The evolution of modern wheat and the relation withAegilops species have been the concern of many scientistsfor many years. Three genomes, namely A, B (S) and D,took part in the evolution of modern wheats. The donorof the A genome is assumed to be T. urartu (Valkoun etal., 1997; Gitte et al., 2006). The donor of the B genome

has also been a matter of discussion among scientists. Itwas claimed by Dvorák (1998) that the B genome isrelated to the genome of Ae. speltoides. Recently, Gitte etal. (2006) declared that the hitherto enigmatic B genomeis derived from Ae. speltoides and the A genome isderived from T. urartu. There is full consensus on thedonor of the D genome, which is Ae. tauschii (Kimber andFeldman, 1987; Waines, 1997; Gitte et al., 2006).

The genus Aegilops (except Aegilops mutica Boiss.)consists of 5 sections: Sitopsis (6 spp.), Aegilops (8 spp.),Vertebrata (5 spp.), Cylindropyrum (2 spp.) andComopyrum (2 spp.) (Van Slageren, 1994).

Kün (1979) agro-morphologically characterized andevaluated some pathological and technological characteristicsof 222 wild wheat species. The study was performed underthe following 5 headings: (1) evolution and nomenclature ofwheat, (2) studies to give male sterility characteristics towheat, (3) role of wild wheat in gaining disease resistance,(4) role of wild wheat in giving quality traits to wheat, and(5) capability of wild wheat in natural vegetation. In thatstudy, it was reported that all populations of Ae. speltoidesvar. speltoides, Ae. speltoides var. ligustica and Ae. biuncialiswere resistant to yellow rust; plant height ranged between26.8 (Ae. umbelleulata) and 97.7 cm (Ae. mutica); spikelength ranged between 27.2 (Ae. umbellulata) and 273.1mm (Ae. mutica); and protein content of the species was upto 32.6% (Ae. umbellulata).

Peflkircio¤lu (1996) characterized and evaluated 13Aegilops and 3 wild Triticum species of Turkish origin fortheir agro-morphological, pathological, and quality traits.Among the material, plant height ranged between 16.6(Ae. juvenalis) and 112.0 cm (Ae. mutica); spike lengthranged between 2.4 (Ae. ovata) and 23.3 cm (Ae.mutica); protein content ranged between 19.9% (Ae.speltoides) and 26.2% (Ae. juvenalis).

Wild relatives of crops play a significant role in thedevelopment of cultivars with desired characteristics.Srivastava and Damania (1989) reported that wildrelatives provide valuable genes for disease resistance,high protein content, tillering, drought resistance andother economically desirable attributes. Lange andBalkema-Boomstra (1988) stated that progenitors ofcultivated species should be considered an importantsource of variability for breeding genetic bases ofcultivated crops. Efficient utilization of these speciesrequires detailed knowledge of their genetic, cytogeneticand agromorphological characteristics.

Agro-Morphological Characterization of Some Wild Wheat (Aegilops L. and Triticum L.) Species

388

The primary aim of this study was to examine theagromorphological characteristics of wild wheat species(Aegilops and Triticum) including the primary relativesthat have a potential to be used in breeding programs.

Materials and Methods

The material consisted of 124 wheat populations, 112of which were collected in marginal areas of southeastand east Turkey. Scientific names of the experimentalmaterial, their genomes, and number and status of thesamples are given in Table 1. Wild materials werecollected during 2 major periods, 1987 (K›z›ltan et al.,1990) and 1994-1998 (Karagöz, 1996), and severalothers. During the first mission, wild wheat material inareas lower than 550 m around Atatürk Dam wascollected. During 1994-1998, almost all of the marginalareas of Ceylanp›nar State Farm, where elevation is 330-550 m, were covered. In the meantime, wild wheatmaterial in north and northeastern parts of Karacada¤Mountain with an elevation range of 800-1600 m werealso collected. Some of the previously collected wildwheat material of Eastern Anatolian origin was includedin the study. Cultivated materials were collected fromremote villages, where landraces have still not been

replaced by modern cultivars. Data on the place of originand altitude of the populations are given in Table 2.

Among the populations tested, Ae. tauschii Coss. var.meyeri (Griseb. ex Ledeb.) samples were collected fromfianlıurfa province. This species of the genus Aegilops hasnot been described in the flora of Turkey and ischaracterized for the first time in this study.

Seeds of the material were germinated in thegreenhouse in September 2001 and transplanted atHaymana-Ankara Research Farm of the Central ResearchInstitute for Field Crops in October 2001. Each plot was3 m long; distance between rows was 35 cm and betweenplants was 20 cm (Delacy et al., 2000: Karagöz andZencirci, 2005; Zencirci and Karagöz, 2005). Plots werenot replicated. The material was characterized for thefollowing 5 major discriminating characters (IBPGR,1978; IBPGR, 1985) during 2002.

Plant height: from ground level to end of spikeexcluding awns at maturity (cm).

Days to heading: number of days when 50% of theplants of each plot headed (Jan. 1 = 1).

Growth habit: 1: upright, 3: semi prostrate, 5:prostrate

A. KARAGÖZ, N. P‹LANALI, T. POLAT

389

Table 1. Scientific name of the experimental material, number and status of the samples.

Scientific name

Aegilops speltoides Tausch var. speltoides

Aegilops speltoides Tausch var. ligustica (Savign.) Fiori.

Aegilops biuncialis Vis.

Aegilops columnaris Zhuk.

Aegilops triuncialis L.

Aegilops neglecta Req. ex Bertol.

Aegilops umbellulata Zhuk.

Aegilops crassa Boiss.

Aegilops tauschii Coss. var. meyeri (Griseb. ex Ledeb.)

Aegilops vavilovii (Zhuk.) Chennav

Triticum monococcum L. subsp. aegilopoides (Link) Thell.

Triticum urartu Tum. ex. Gand.

Triticum turgidum L. subsp. dicoccoides (Körn. ex Asch. & Graebn.)

Triticum turgidum L. subsp. dicoccon (Schrank) Thell.

Triticum monococcum L. subsp. monococcum

Abbreviation

Ae. speltoides

Ae. ligustica

Ae. biuncialis

Ae. columnaris

Ae. triuncialis

Ae. neglecta

Ae. umbellulata

Ae. crassa

Ae. tauschii

Ae. vavilovii

T. aegilopoides

T. urartu

T. dicoccoides

T. dicoccon

T. monococcum

Genome

S

S

UM

UM

UC

UM, UMUn

U

DM

D

DMS

A

A

AB

AB

A

No. ofpopulations

14

9

11

16

22

2

14

2

2

2

3

2

13

8

4

Section ofAegilops spp.

SitopsisSitopsis

Aegilops

Aegilops

Aegilops

Aegilops

Aegilops

Vertabrata

Vertabrata

Vertabrata

-

-

-

-

-

Status

Wild

Wild

Wild

Wild

Wild

Wild

Wild

Wild

Wild

Wild

Wild

Wild

Wild

Cultivated

Cultivated


390

No. Pop. No. Species Province Alt. (m)

1 TUR 00047 Ae. biuncialis Ad›yaman 4102 TUR 00051 Ae. biuncialis Ad›yaman 4103 TUR 00102 Ae. biuncialis Ad›yaman 4004 TUR 00307 Ae. biuncialis Ad›yaman 5855 TUR 00385 Ae. biuncialis Ad›yaman 4556 TUR 00407 Ae. biuncialis Ad›yaman 4707 TUR 00483 Ae. biuncialis Ad›yaman 4508 TUR 00633 Ae. biuncialis Ad›yaman 5909 TUR 00788 Ae. biuncialis fianl›urfa 47510 TUR 01406 Ae. biuncialis fianl›urfa 52511 TUR 01615 Ae. biuncialis fianl›urfa 45012 TUR 00015 Ae. columnaris Ad›yaman 41013 TUR 00053 Ae. columnaris Ad›yaman 41014 TUR 00110 Ae. columnaris Ad›yaman 63015 TUR 00211 Ae. columnaris Ad›yaman 41516 TUR 00227 Ae. columnaris Ad›yaman 54017 TUR 00319 Ae. columnaris Ad›yaman 58518 TUR 00674 Ae. columnaris Ad›yaman 65519 TUR 00772 Ae. columnaris fianl›urfa 47520 TUR 00773 Ae. columnaris fianl›urfa 48521 TUR 00794 Ae. columnaris fianl›urfa 45522 TUR 00814 Ae. columnaris fianl›urfa 58023 TUR 00936 Ae. columnaris fianl›urfa 48524 TUR 01000 Ae. columnaris fianl›urfa 52525 TUR 01134 Ae. columnaris fianl›urfa 47526 TUR 01178 Ae. columnaris fianl›urfa 55027 TUR 00358 Ae. columnaris Ad›yaman 48028 TUR 01161 Ae. crassa fianl›urfa 47529 TUR 02605 Ae. crassa fianl›urfa 48030 TUR 02554 Ae. tauschii fianl›urfa 48031 TUR 02770 Ae. tauschii fianl›urfa 40032 TUR 00002 Ae. triuncialis Ad›yaman 41033 TUR 00103 Ae. triuncialis Ad›yaman 40034 TUR 00252 Ae. triuncialis Ad›yaman 53035 TUR 00343 Ae. triuncialis Ad›yaman 51536 TUR 00366 Ae. triuncialis Ad›yaman 48037 TUR 00402 Ae. triuncialis Ad›yaman 47038 TUR 00430 Ae. triuncialis Ad›yaman 59539 TUR 00454 Ae. triuncialis Ad›yaman 54040 TUR 00544 Ae. triuncialis Ad›yaman 53041 TUR 00569 Ae. triuncialis Ad›yaman 57542 TUR 00632 Ae. triuncialis Ad›yaman 59043 TUR 00728 Ae. triuncialis Ad›yaman 47544 TUR 00879 Ae. triuncialis fianl›urfa 55045 TUR 01037 Ae. triuncialis fianl›urfa 51046 TUR 01098 Ae. triuncialis fianl›urfa 47547 TUR 01177 Ae. triuncialis fianl›urfa 55048 TUR 01257 Ae. triuncialis fianl›urfa 52549 TUR 01396 Ae. triuncialis fianl›urfa 57550 TUR 01407 Ae. triuncialis fianl›urfa 52551 TUR 01687 Ae. triuncialis fianl›urfa 55052 TUR 01776 Ae. triuncialis fianl›urfa 82553 TUR 01778 Ae. triuncialis fianl›urfa 85054 TUR 00066 Ae. neglecta Ad›yaman 40055 TUR 00983 Ae. neglecta fianl›urfa 44556 TUR 00223 Ae. umbellulata Ad›yaman 54057 TUR 00224 Ae. umbellulata Ad›yaman 55058 TUR 00270 Ae. umbellulata Ad›yaman 55559 TUR 00355 Ae. umbellulata Ad›yaman 51560 TUR 00417 Ae. umbellulata Ad›yaman 59561 TUR 00708 Ae. umbellulata Ad›yaman 83062 TUR 00729 Ae. umbellulata Ad›yaman 475

No. Pop. No. Species Province Alt. (m)

63 TUR 01036 Ae. umbellulata fianl›urfa 51064 TUR 01072 Ae. umbellulata fianl›urfa 47565 TUR 01577 Ae. umbellulata fianl›urfa 56066 TUR 01769 Ae. umbellulata Ad›yaman 80067 TUR 00787 Ae. umbellulata Ad›yaman 47568 TUR 02220 Ae. umbellulata Erzincan 111069 TUR 02632 Ae. umbellulata fianl›urfa 40070 TUR 01214 Ae. vavilovii fianl›urfa 50071 TUR 01248 Ae. vavilovii fianl›urfa 52572 TUR 00488 Ae. speltoides Ad›yaman 46073 TUR 00623 Ae. speltoides Ad›yaman 62074 TUR 00634 Ae. speltoides Ad›yaman 59075 TUR 00903 Ae. speltoides fianl›urfa 57576 TUR 01523 Ae. speltoides fianl›urfa 52577 TUR 01636 Ae. speltoides fianl›urfa 48578 TUR 01690 Ae. speltoides fianl›urfa 55079 TUR 01725 Ae. speltoides fianl›urfa 60080 TUR 01765 Ae. speltoides fianl›urfa 80081 TUR 02556 Ae. speltoides fianl›urfa 45082 TUR 02764 Ae. speltoides Adana 110083 TUR 02785 Ae. speltoides Gaziantep 85084 TUR 03425 Ae. speltoides Hatay 90085 TUR 03499 Ae. speltoides fianl›urfa 48086 TUR 00301 Ae. ligustica Ad›yaman 58587 TUR 02210 Ae. ligustica Erzincan 118088 TUR 02774 Ae. ligustica fianl›urfa 45089 TUR 03387 Ae. ligustica Diyarbak›r 80090 TUR 03352 Ae. ligustica Diyarbak›r 75091 TUR 03354 Ae. ligustica Diyarbak›r 77592 TUR 03355 Ae. ligustica Diyarbak›r 79093 TUR 03374 Ae. ligustica Diyarbak›r 80094 TUR 03384 Ae. ligustica Diyarbak›r 80095 TUR 02440 T. dicoccon Sinop 110096 TUR 02453 T. dicoccon Sinop 120097 TUR 02456 T. dicoccon Sinop 100098 TUR 03558 T. dicoccon Sinop 110099 TUR 03560 T. dicoccon Kastamonu 1200100 TUR 03562 T. dicoccon Kastamonu 1200101 TUR 03564 T. dicoccon Kastamonu 1100102 TUR 03565 T. dicoccon Kastamonu 1050103 TUR 00842 T. dicoccoides fianl›urfa 580104 TUR 02637 T. dicoccoides fianl›urfa 480105 TUR 03346 T. dicoccoides Diyarbak›r 750106 TUR 03358 T. dicoccoides Diyarbak›r 725107 TUR 03362 T. dicoccoides Diyarbak›r 750108 TUR 03369 T. dicoccoides Diyarbak›r 740109 TUR 03371 T. dicoccoides Diyarbak›r 740110 TUR 03376 T. dicoccoides Diyarbak›r 750111 TUR 03388 T. dicoccoides Diyarbak›r 700112 TUR 03391 T. dicoccoides Diyarbak›r 700113 TUR 03396 T. dicoccoides Diyarbak›r 700114 TUR 03399 T. dicoccoides Diyarbak›r 690115 TUR 03402 T. dicoccoides Diyarbak›r 690116 TUR 00241 T. aegilopoides Ad›yaman 650117 TUR 00741 T. aegilopoides Ad›yaman 570118 TUR 00803 T. aegilopoides fianl›urfa 530119 TUR 02412 T. monococcum Sinop 800120 TUR 03559 T. monococcum Karabük 1020121 TUR 03561 T. monococcum Kastamonu 1150122 TUR 03563 T. monococcum Karabük 1020123 TUR 00734 T. urartu Ad›yaman 570124 TUR 00853 T. uratru fianl›urfa 590

Table 2. Data on place of origin and altitude of the populations.

Number of stems: counted at maturity

Spike length: length of spikes excluding awns atmaturity (mm).

Observations were performed on 5 randomly takenplants from each population. Variety specificcharacteristics such as spike density, and awnedness werenot taken into consideration since there was no variationin such characteristics. Likewise, days to maturity traitwas also discarded because of its high similarity with daysto heading.

Mean, standard deviation (SD), coefficient of variation(CV), and range were computed for each population bymeans of the Tarist statistical program; and clusteranalysis based on a Euclidian similarity matrix wascomputed on average values of populations by means ofthe Minitab statistical program. The similaritydendrogram for the species is given in Figure 1.

Results and Discussion

Depending on the size of samples, noticeable variationwas found for plant height, growth habit, number ofstems, and spike length. The observed variation impliesthat populations could provide a variety of genotypes forfurther studies. Very high CV values have been obtainedin some of the characteristics such as growth habit,number of stems, and spike length. Dotlacil et al. (2000)considered a minimum 10% CV a sign of wide diversity inwheat landraces and obsolete varieties. Values obtained ineach trait are given below:

Plant height:

Plant height is an important characteristic in wheatbreeding. Modern high yielding wheat cultivars areshorter than landraces and old cultivars. A wide range ofvariation was observed in Ae. speltoides, Ae. columnarisand Ae. umbellulata species, with CV values above 10%


391

T. aegilopoides

T. urartu

T. monococcum

T. dicoccoides

T. dicoccon

Ae. ligustica

Ae. aucheri

Ae. vavilovü

Ae. tauschü

Ae. crassa

Ae. triuncialis

Ae. columnaris

Ae. neglecta

Ae. umbellulata

Ae. biuncialis

64 76 88 100

Triticum

Sitopsis

Vertebrata

Aegilops

Figure 1. Cluster analysis based on a Euclidian similarity matrix for the species.

(Table 3). The mean values given by Kün (1979) are afew centimeters shorter than our figures, and totallyharmonious in Ae. ligustica. Our figures are compatiblewith those given by Peflkircio¤lu (1996) for Ae. triuncialisand Ae. umbellulata, and comparable for A. columnaris,Ae. biuncialis and Ae. neglecta. Our values for T.aegilopoides, T. urartu and T. dicoccoides are higher thanPeflkircio¤lu’s (1996) findings and generally shorter thanVan Slageren’s (1990) figures except for Ae. speltoidespopulations. Differences in plant height figures aresupposed to have been caused by the origin of thesamples.

Among the T. dicoccoides samples, fianlıurfa originmaterial formed a separate cluster, with lower plantheight values. Wild wheat material taken from lowerelevations of Karacada¤ is considerably taller than theothers. Damania (1993) reported a similar trend in T.dicoccoides, i.e. robust early maturing types are growingin warmer habitats and more slender later bloomingtypes at higher elevations.

Our findings are in harmony with those reported byHumeid et al. (1988) for Aegilops species. In themeantime, Turkish origin T. dicoccoides samples tested inSyria were 23.0 cm shorter than those in Turkey. Thelack of information about the collecting sites does notallow us to make any meaningful comment about this.

Extreme values for this trait were observed for Ae.umbellulata and T. urartu (Table 3). Among the Aegilopsspecies, the highest variation was observed in Ae.umbellulata. Regarding the Triticum species, wildmaterial was taller than the cultivated material (Table 3).Mean values for this trait were the smallest for thesection Aegilops, followed by Vertebrata and Sitopsis.Very close values were observed in Sitopsis material forthis trait (Table 4).

Days to heading:

Under favorable conditions, a longer vegetationperiod suggested higher yield, but, in areas like CentralAnatolia where a dry period is inevitable after spring,earliness of the cultivated material is becoming moreimportant. The lowest SD and CV values were detectedfor this trait (Table 3). This implies that variation amongthe population was not large enough for days to heading.

Noticeable populations for this trait were TUR 03391(T. dicoccoides) (175 days), TUR 01636 (187 days) andTUR 01690 (Ae. speltoides) (187 days), and TUR 00366

and TUR 01687 (Ae. triuncialis) (190 days) for lateheading; and TUR 00842 and TUR 02637 (T.dicoccoides) (155 days) for early heading

Vertebrata species were the earliest ones, followed byAegilops and Sitopsis. Wild Triticum species were a fewdays earlier than the cultivated ones (Table 3). Ae. crassawas the earliest species, followed by Ae. tauschii. Tallgrowing species seemed to have headed latest of all. Thehighest variation was observed in Ae. speltoides.

Heading time of all the species is totally compatiblewith those reported by Kün (1979) and Peflkircio¤lu(1996). A larger range was observed in Ae. triuncialis,Ae. speltoides and Ae. columnaris (Table 3). A similartrend was reported for Ae. triuncialis and Ae. umbellulataby Kün (1979). In comparison with the results of wildwheat evaluation studies conducted in Syria, Ae. tauschiisamples were 2 days earlier in Ankara conditions, whileAe. columnaris, Ae. vavilovii, Ae. biuncialis, Ae. neglectaand Ae. triuncialis samples were 10.6-30.6 days later forheading (Van Slageren and Sweid, 1991). Contrastingresults have been reported by Humeid et al. (1998) andVan Slageren and Sweid (1991). Ae. triuncialis wasreported to be the earliest heading by Van Slageren andSweid (1991), while it was found to be the latest headingone by Humeid et al. (1988). Figures reported by vanSlageren and Sweid (1991) are harmonious with ourfindings only in terms of order of heading time byspecies.

Among the Ae. triuncialis population samples, extremevalues for this trait were observed in TUR 01396 (139days) and TUR 01687 (190 days). Both of thepopulations were collected from nearby habitats atalmost the same altitudes. Remarkable variations amongthe neighboring populations suggest significant geneticdifferences between them.

Growth habit:

Among the characters observed, the highest variationwas observed in growth habit, with 0.00% to 55.98%CV values. All the Triticum species excluding T.dicoccoides were upright. Ae. ligustica was almostupright, while Ae. speltoides was near semiprostrate. Ae.tauschii was prostrate. The rest of the Aegilops materialranked between near semiprostrate (Ae. triuncialis) andnear prostrate (Ae. biuncialis) (Table 3).

Ae. columnaris, Ae. ligustica and T. dicoccoidesshowed a high degree of variation for this characteristic,


392


393

Table 3. Mean, minimum, maximum, SD and CV for plant height, days to heading, growth habit, number of stemsand spike length of the species.

Species Plant Days to Growth Number Spikeheight heading habit of stems length

Mean 77.23 160.41 2.26 68.96 128.81

SD 2.15 3.27 0.34 3.97 6.38

Ae. speltoides CV 10.40 7.63 55.98 21.56 18.53

Min. 46 144 1 22 72

Max. 105 187 5 170 185

Mean 76.31 152.07 1.47 60.18 95.22

SD 1.80 1.98 0.20 8.69 3.26

Ae. ligustica CV 7.08 3.91 40.91 43.33 10.28

Min. 64 140 1 12 67

Max 105 161 3 203 145

Mean 39.07 148.47 4.18 58.24 34.58

SD 1.08 1.60 0.35 2.97 1.08

Ae. biuncialis CV 9.15 3.58 2.92 16.94 10.32

Min. 27 143 3 30 23

Max 55 158 5 102 45

Mean 41.49 153.44 3.93 74.01 43.40

SD 1.28 2.02 0.30 4.57 4.24

Ae. columnaris CV 12.29 5.27 40.24 24.71 39.08

Min. 24 140 1 27 25

Max 54 175 5 198 111

Mean 37.00 158.90 3.00 47.40 31.40

SD 1.40 1.90 0.00 0.40 0.80

Ae. neglecta CV 5.35 1.69 0.00 1.19 3.60

Min. 22 157 3 21 26

Max. 41 174 3 67 37

Mean 44.92 154.27 3.42 68.23 58.16

SD 0.96 1.96 0.17 5.55 2.28

Ae. triuncialis CV 9.98 5.96 23.54 38.18 18.36

Min. 30 139 3 15 25

Max 53 190 5 175 79

Mean 30.49 151.86 3.83 59.69 28.47

SD 8.89 1.72 0.32 5.13 1.45

Ae. umbellulata CV 10.97 4.24 30.79 32.14 19.03

Min. 21 140 3 21 18

Max 54 159 5 133 45

Mean 41.90 137.20 3.00 10.60 67.10

SD 1.10 0.40 0.00 0.40 0.10

Ae. crassa CV 3.71 0.41 0.00 5.34 0.21

Min. 32 135 3 7 54

Max 50 138 3 15 79


394

Table 3. Mean, minimum, maximum, SD and CV for plant height, days to heading, growth habit, number of stemsand spike length of the species (continued).

Species Plant Days to Growth Number Spikeheight heading habit of stems length

Mean 38.10 140.00 5.00 28.50 75.50

SD 2.10 0.00 0.00 8.50 1.50

Ae. tauschii CV 7.79 0.00 0.00 42.18 2.81

Min. 31 140 5 12 64

Max 62 140 5 73 88

Mean 49.80 142.40 3.00 24.50 110.00

SD 0.60 1.20 0.00 6.30 4.40

Ae. vavilovii CV 1.70 1.19 0.00 36.37 5.66

Min. 47 140 3 12 66

Max 55 144 3 78 135

Mean 109.40 159.33 1.00 27.67 133.60

SD 6.16 1.38 0.00 2.71 9.53

T. aegilopoides CV 9.75 1.50 0.00 16.96 12.35

Min. 43 155 1 11 93

Max 132 162 1 67 173

Mean 119.70 159.60 1.00 25.00 103.50

SD 3.50 0.60 0.00 5.80 4.90

T. urartu CV 4.14 0.53 0.00 32.81 6.70

Min. 103 151 1 17 95

Max 134 165 1 35 115

Mean 88.68 162.02 1.92 21.74 96.68

SD 2.36 1.01 0.23 1.24 3.40

T. dicoccoides CV 9.61 2.24 42.78 20.51 12.70

Min. 50 155 1 7 55

Max 110 175 3 55 142

Mean 98.35 165.65 1.00 39.50 86.70

SD 7.26 1.05 0.00 2.77 5.21

T. monococcum CV 14.77 1.26 0.00 14.03 12.02

Min. 64 162 1 13 65

Max 113 169 1 79 98

Mean 81.70 163.90 1.00 29.85 73.23

SD 3.76 1.08 0.00 1.58 2.33

T. dicoccon CV 13.03 1.86 0.00 15.00 9.01

Min. 44 161 1 10 61

Max 112 170 1 65 90


395

Tabl

e 4.

Mea

n, S

D a

nd C

V va

lues

for

Aeg

ilops

and

Triti

cum

grou

ps.

Plan

t H

eigh

tD

ays

to h

eadi

ngG

row

th h

abit

No.

of

stem

sSp

ike

leng

th

Mea

nSD

CVM

ean

SDCV

Mea

nSD

CVM

ean

SDCV

Mea

nSD

CV

Sito

psis

76.7

77.

19.

1115

6.24

11.0

47.

031.

861.

1257

.55

64.5

719

.93

30.4

111

2.02

25.5

422

.08

Aegi

lops

grou

psAe

gilo

ps38

.59

6.0

16.7

815

3.39

7.80

5.12

3.49

1.21

31.7

861

.51

32.0

932

.09

39.2

015

.98

36.5

1

Vert

ebra

ta43

.27

7.21

15.7

813

9.87

3.41

2.41

3.67

1.03

28.1

720

.70

40.4

240

.42

84.4

624

.30

30.3

7

Triti

cum

grou

psW

ild10

5.93

9.85

9.85

160.

322.

451.

531.

310.

8725

.74

30.4

722

.71

67.2

511

1.26

18.4

514

.49

Culti

vate

d90

.03

13.0

315

.36

164.

783.

812.

311.

000.

0000

.00

34.6

834

.68

18.6

079

.96

9.56

12.4

4

with CV values over 40% (Table 3). Regarding thecollection sites, Ae. columnaris (TUR 00110) and Ae.speltoides (TUR 03499) samples were the most variableones, with this character ranking from 1 to 5. fianlıurfaorigin materials were generally semiprostrate. Significantvariation was also observed in T. dicoccoides samples. Allof the cultivated Triticum species were upright, while wildones were close to upright. Among the Aegilops species,the section Sitopsis was close to upright while thesections Aegilops and Vertebrata were almostsemiprostrate (Table 4).

The materials collected on Karacada¤ Mountain inDiyarbakır province were the most variable ones, rankingbetween 1 and 3. This is mainly due to the altitudes of thecollecting sites. Material from lower elevation areastended to be more upright. Our observations arecompatible with those reported by Damania (1993) inthis regard. High degree of CV values and considerabledifferences between the populations collected fromrelatively nearby areas are an indication of geneticdiversity between populations. Therefore, in areas withvery high genetic diversity it is worth collecting from veryclose distances. Our findings are compatible with thosepublished by Peflkircio¤lu (1996), except for minordifferences caused by the index used for scoring.

Number of stems:

This is rather an important characteristic forsustainability of species in the wild, since the number ofstems is related to the number of spikes and consequentlythe number of seeds produced by each species. Amongthe accessions, the most remarkable ones were TUR01636 (Ae. speltoides), TUR 00301 and TUR 02210 (Ae.ligustica), and TUR 01776 (Ae. triuncialis) for their hightillering capacity; and TUR 00307 (Ae. biuncialis) andTUR 00053 (Ae. columnaris) for their low number oftillers. T. dicoccoides populations TUR 00842 and TUR02637 also showed higher tillering capacity than theothers.

There was very high degree of variation in this traitfor all species except Ae. crassa and Ae. neglecta (Table3). It is generally the case that members of the sectionVertebrata are not abundant in nature and they are alsoa difficult section for collecting sufficient amount ofseeds, because of the scarcity of the stems they produce.What is observed in situ has also been reflected in fieldconditions. Vertebrata species were the least productive

group in this regard, followed by wild and cultivatedTriticum species. The most productive Aegilops sectionswere Aegilops and Sitopsis (Table 4).

Our findings were quite compatible with thosereported by Peflkircio¤lu (1996) for section Aegilops andsection Vertebrata species. Considerable differences wererecorded for section Sitopsis and wild Triticum speciesbetween our findings and the references cited. Ourfigures were significantly higher than the values reportedby Humeid et al. (1998) and Van Slageren and Sweid(1991). It is thought that the main difference is caused bythe higher tillering capacity of Turkish material.

Spike length:

High variation was observed in all the species exceptfor Ae. neglecta, Ae. crassa, Ae. vavilovii, Ae. tauschii andT. urartu. Extreme values were obtained in Ae.umbellulata, Ae. columnaris and Ae. speltoides species forthis character. While Ae. neglecta and Ae. biuncialis wereshorter than average, Ae. vavilovii and Ae. ligustica werethe tallest heading ones (Table 3).

Among the Aegilops sections and Triticum groups, thesection Sitopsis gave the highest values, followed by wildTriticum species. The section Aegilops gave the minimumaverage value. Cultivated Triticum group and sectionVertebrata were ranked in between (Table 4).

Our findings were compatible with the figuresreported by Kün (1979) and Peflkircio¤lu (1996) for Ae.ligustica, Ae. biuncialis, Ae. triuncialis and Ae. neglecta;and compatible with those given by Peflkircio¤lu (1996)for A. columnaris, Ae. umbellulata and T. urartu. In therest of the material we observed higher values for all butAe. crassa. The insufficient number of samples for Ae.crassa does not allow us to make a meaningful commentabout it.

Cluster analysis based on a Euclidian similarity matrixfor 5 characters ranked the species meaningfully (Figure1). The dendrogram is compatible with diagrammaticrepresentation sectional organization of Aegilops species(Van Slageren, 1994). Species of the section Aegilopswere in the first cluster, followed by the sectionsVertebrata and Sitopsis. The last cluster was formed byTriticum species. The highest similarity was observedbetween Ae. biuncialis and Ae. umbellulata (91.6%). Veryhigh similarity values were detected between Ae.biuncialis and Ae. triuncialis (88.3%), Ae. columnaris andAe. triuncialis (87.7%), Ae. crassa and Ae. tauschii


396

(85.5%), T. dicoccon and T. dicoccoides (82.2%), andAe. speltoides and Ae. ligustica (71.7%).

As a result of this study, we can conclude that therewas significant agro-morphological variation between thepopulations for the traits studied. The high degree ofvariation found in the material collected along shortdistances suggests that ex-situ conservation programsshould take into consideration that fact by sampling overshort distances in order to capture as much of the

variation as possible. Samples from certain parts offianlıurfa and Adıyaman provinces as well as the whole ofKaracada¤ Mountain and nearby areas should be collectedand characterized in a systematic manner.

Acknowledgments

We sincerely appreciate the help of our colleaguesGülay Sa¤lam and Hale Sümen during the field research.


397

References

Damania, A.B. 1993. Domestication and in situ conservation of cerealwild progenitors in the Near East. In: Biodiversity and WheatImprovement (Ed.: A.B. Damania). Wiley.

Davis, P.H. 1985. Flora of Turkey and the East Aegean Islands. V:9.Edinburgh University Press, Edinburgh.

Delacy, I.H., B. Skowmand and J. Huerta. 2000. Characterization ofMexican landraces using agronomically useful attributes. GeneticResources and Crop Evolution. 47: 87-96.

Diamond, J. 1997. Location, location, location: The first farmers.Science, 278: 1243-1244.

Dotlacil, L., J. Hermuth, Z. Stehno and M. Manev. 2000. Diversity inEuropean winter wheat landraces and obsolete cultivars. Czech J.Genet. Plant Breeding, 16: 29-36.

Dvorák, J. 1998. Genome analysis in the Triticum-Aegilops alliance. In:Proc. of the 9th Int. Wheat Genetics Symp. (Ed.: A.E. Slinkard).Vol. 1., 8-11. Univ. of Saskatchewan.

Gitte, P., O. Seberg, M. Yde and K. Berthelsen. 2006. Phylogeneticrelationships of Triticum and Aegilops and evidence for the originof the A, B, and D genomes of common wheat (Triticumaestivum). Molecular Phylogenetics and Evolution, 39: 70-82.

Gökgöl, M. 1935. Türkiye Bu¤dayları. Die Türkischen Weizen. V. 1.‹stanbul.

Harlan, J.R. 1981. The early history of wheat: Earliest traces to thesack of Rome. In: Wheat Science – Today and Tomorrow. (Eds.:L.T. Evans and W.J. Peacock). Cambridge Univ. Press,Cambridge, UK.

Heun, M., R. Schafer-Pregl, D. Klawan, R. Castagna, M. Accerbi, B.Borghi and F. Salamini. 1997. Site of Einkorn wheatdomestication identified by DNA fingerprinting. Science, 278:1321-1314.

Humeid, B., O.F. Mamluk and A.A. Jaradat. 1988. Annual Report for1988, Genetic Resources Unit. ICARDA. 21-23. Aleppo.

IBPGR. 1978. Descriptors for Wheat & Aegilops. A Minimum List.IBPGR Secreteriat, Rome.

IBPGR. 1985. Descriptors of Wheat (revised). IBPGR Secretariat,Rome.

Karagöz, A. 1996, In situ conservation of plant genetic resources inCeylanpınar State Farm. In: The Proc. of the Int. Symp. on In situConservation of Plant Genetic Diversity. (Eds.: N. Zencirci, Z.Kaya, Y. Anikster and W.T. Adams). P 87-92, Ankara.

Karagöz, A. and N. Zencirci. 2005. Variation in wheat (Triticum spp.)landraces from different altitudes of three regions of Turkey.Genetic Res. Crop Evolution. 52: 775-785.

Kızıltan, M., A. Karagöz and J.G. Waines. 1990. The plant expeditionresearch from south eastern Anatolia (Atatürk Dam) C7 square.Wheat Information Service, 70: 15-28.

Kimber, G. and M. Feldman. 1987. Wild Wheat. An Introduction. SpecialReport 353, College of Agriculture University of Missouri-Columbia.

Kün, E. 1979. Türkiye Yabani Bu¤daylarının (Aegilops L.) IslahYönünden Önemli Bazı Karakterleri, Üzerinde Arafltırmalar. A.Ü.Ziraat Fakültesi Yayın No: 685, Bilimsel Arafltırma ve ‹ncelemeler:400, Ankara.

Lange, W. and A.G. Balkema-Boomstra. 1988. The use of wild speciesin breeding barley and wheat, with specific reference to theprogenitors of the cultivated species. In: Cereal Breeding Relatedto Integrated Cereal Production. Eucarpia. (Eds.: M.L. Joena andL.A.J. Slootmaker). Wageningen, The Netherlands.

Lev-Yadun, A., A. Gopher and S. Abbo. 2000. The cradle of agriculture.Sci. 288; 1602-1603.

Nesbit, M. and L. Samuel. 1998. Wheat domestication, archeobotanicalevidence. Science. 279: 1433.

Nevo, E., A.B. Korol, A. Beiles and T. Fahima. 2002. Evolution of WildEmmer and Wheat Improvement. Population Genetics, GeneticResources and Genome Organization of Wheat’s Progenitor,Triticum dicoccoides. Springer, Heidelberg.

Peflkircio¤lu, M.H. 1996. Türkiye’de Yetiflen Yabani Bu¤day Türlerinin(Triticum spp. ve Aegilops spp.) Bazı ‹leri Tanımlama Özellikleri.Doktora Tezi. A. Ü. Fen Bilimleri Enstitüsü. p 185.

Srivastava, J.P. and A.B. Damania. 1989. Use of collections in cerealimprovement in semi-arid areas. In: The Use of Plant GeneticResources. (Eds.: A.H.D. Brown et al.). Cambridge UniversityPress, Cambridge, UK.


398

Valkoun, J., J.G. Waines, and J. Konopka. 1997. Current GeographicalDistribution and Habitat of Wild Wheats and Barley. In: Proc. ofHarlan Symposium. 10-14 May Aleppo. (Eds.: Damania et al.).Part 6. www.ipgri.cgiar.org/Publications/ HTMLPublications/

Van Slageren, M.W. 1994. Annual Report for 1991, Genetic ResourcesUnit. ICARDA. 46-48. Aleppo.

Van Slageren, M.W. 1994. Wild wheats: a monograph of Aegilops L. andAmblyopyrum (Jaub. & Spach) Eig. (Poaceae). WageningenAgricultural University, ICARDA.

Waines, J.G. 1997. In situ conservation of wild relatives of crop plantsin relation to their history. Conservation of wild progenitors. In:Proc. of Harlan Symposium. 10-14 May Aleppo. (Eds.: Damaniaet al.). Part 6. ww.ipgri.cgiar.org/Publications/HTMLPublications/

Zencirci, N. and A. Karagöz. 2005. Effect of developmental stageslength on yield and some quality traits of Turkish durum wheat(Triticum turgidum L. convar. durum (Desf.) Mackey) landraces.Genetic Res. Crop Evolution. 52: 765-774.

Agro-Morphological Characterization of Some Wild Wheat (Aegilops L. and Triticum L.) Species

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