ENGLISH TRANSLATION OF THE 1979 RUSSIAN TAXONOMIC MONOGRAPH OF TRITICUM L.
BY DOROFEEV ET AL.: PROJECT PROGRESS REPORT

 

Helmut Knüpffer1, Laura A. Morrison2, Anna A. Filatenko3, Karl Hammer4, Alexei Morgounov5, Iva Faberová6

1Genebank, Institute of Plant Genetics and Crop Plant Research (IPK), D-06466 Gatersleben (Germany)
Phone +49 39482 5283, Fax +49 39482 5155, E-mail knupffer@ipk-gatersleben.de

2Department of Crop and Soil Science, Oregon State University, Corvallis, OR 97331-3002 (USA)

313-Linija 12, kv. 7, St. Petersburg 199 034 (Russia)

4Universität Gesamthochschule Kassel, Steinstraße 11, D-37213 Witzenhausen (Germany)

5CIMMYT, P.O. Box 374, Almaty 480000 (Kazakhstan)

6Genebank, Research Institute of Crop Production, Drnovská 507, CZ-161 06 Prague (Czech Republic)

 

 

ABSTRACT: Dorofeev et al. (1979), the taxonomic monograph of Triticum L. published by the N. I. Vavilov Institute of Plant Industry (VIR) is an important work which is largely unknown outside of Russia due to the language barrier. A morphological treatment, this monograph is at odds with current genetic concepts of wheat taxonomy and is therefore controversial. It is the only complete modern catalogue of wild and domesticated forms of Triticum species. For germplasm diversity and intellectual property rights, Dorofeev et al. can prove its value as an authoritative source that can be consulted for issues relating to the validity of proprietary claims on wheat genes and genetic lines that rightfully belong within the public domain. In 1999, an international collaboration was formed to support the translation of Dorofeev et al. into English. A project fund that was started by a $5000 donation from CIMMYT has been established at the Society for the Support of Research on Cultivated Plants in Gatersleben (GFK). The project is currently seeking additional matching funds. Dr. Irina Sokolova of the Komarov Botanical Institute has completed the text translation. Dr. Anna Filatenko, one of the original authors, is currently reviewing the translated copy, and adding necessary details and corrections. English editing will be underway in 2002. Publication of an affordable translation is projected for 2003. Internet copies of the taxonomic keys and descriptions also are planned.

 

 

  1. INTRODUCTION

The Russian taxonomic monograph of Triticum L. by Dorofeev et al. (1), published by the N. I. Vavilov Institute of Plant Industry (VIR) in 1979 and known in Russia as the "Wheat Flora", is an important modern taxonomic treatment that is not widely known. It represents the result of a cumulative effort that began in the early 20th century under the guidance of Vavilov. This monograph stands today as a significant research contribution that is unrecognised for its value by most of the non-Russian speaking wheat research community. The only equivalent work in English is the 1921 monograph of the wheats by John Percival, The Wheat Plant (2), a volume that predates the modern genetic concept of the wheats.

Since its publication, Dorofeev et al. has been unavailable to the majority of the wheat research community due to the Russian language barrier. As a traditional treatment largely based in morphology, this obscure work has been eclipsed by the genetic concepts of Triticum and Aegilops L. Since Bowden’s so-called phylogenetic treatment of the wheats published in 1959 (3), wheat taxonomy has been in the hands of geneticists. While many important genetic contributions on the evolutionary relationships within Triticum and Aegilops have been offered in the past half century, the taxonomic handling of the wheats has suffered from competing genetic concepts and an acrimonious debate over circumscription of the wheat genera and their species.

In actuality, the differences between Dorofeev et al. and the genetically based classifications, which are represented as more phylogenetically accurate, are largely differences of nomenclature. Very little in the overall concept of Triticum has changed since Schulz organised the genus into the morphological groups of einkorn, emmer, and dinkel wheats (4), which proved later to follow the ascending polyploid series that traces the evolutionary development of Triticum species. Dorofeev et al. is no less modern than the classifications of the Bowden school. In fact, the differences are not in phylogenetic treatment but in the ranking and handling of the domesticated wheats. In Dorofeev et al., they are given specific status and are fully described; in the Bowden-type classifications, they are named as infraspecific taxa but poorly described (5), or they are eliminated entirely and thereby lost (6). The prevailing genetic classification concepts have distracted attention away from infraspecific diversity that is the basis for species variation. If infraspecific forms are not named and described systematically in the tradition of "modern" taxonomy, their diversity cannot be exploited and is at risk of being lost.

Many researchers, particularly younger members of the wheat research community who are unaware of the underlying history, have recognised the need to bring order into the chaos that Triticum taxonomy has become. One effort is the GrainTax project which provides updated on-line information about taxonomic classifications and wheat synonymy (http://wheat.pw.usda.gov/ ggpages/GrainTax/description.html) (7, 8). Another such effort is the Dorofeev et al. Translation Project (DTP) which was undertaken in 1999 as an internationally based volunteer project to produce an English-language version of the 1979 Dorofeev et al. Triticum monograph (9).

 

2. IMPORTANCE OF THE TRANSLATION OF DOROFEEV ET AL.

There are several pressing reasons for re-examining the importance of Dorofeev et al.

  1. This monograph is the only modern catalogue of wild and domesticated wheat species and their infraspecific diversity.
  2. As a work built on a long historical research base, it represents a compilation of information that would be extremely difficult to reproduce in today’s research funding climate.
  3. A detailed classification provides a guide for dealing with the tremendous diversity of the genus Triticum existing in natural and man-made habitats as well as preserved in genebanks. It is essential to catalogue international genebank collections with a standardised classification system such as found in Dorofeev et al.

 

3. BENEFITS OF THE TRANSLATION

Although wheat geneticists have relegated traditional taxonomic practice to a relatively minor role, taxonomy now promises to take on a new status as an authoritative resource for establishing the boundaries for protection of germplasm diversity and intellectual property rights. In the rapidly developing markets of biotechnology, the potential threats on the public ownership of wild and domesticated wheat diversity can be challenged with the documentation already present in the comprehensive treatment that Dorofeev et al. embodies. Its detailed morphological classification of domesticated landraces and pre-modern cultivars will be useful in establishing the extent of diversity across the geographic range of wheat distribution, particularly for Asia, Eastern Europe, and North Africa. Dorofeev et al. also will be useful in all aspects of biodiversity and plant genetic resources research, including preservation, cataloguing, and utilisation.

Dorofeev et al. is the only modern catalogue of all known infraspecific taxa of the domesticated and wild wheat species. It serves as an authoritative reference that can be consulted for identification of distinct forms of the wild and domesticated wheats and for establishing public ownership of wheat genes or genetic forms in the face of proprietary ownership challenges by private companies.

 

4. INFORMATIONAL VALUE OF THE MONOGRAPH

The monograph describes 27 Triticum species (cf. Table 1). By contrast, genetic classifications usually recognise only four or five species. While the appropriateness of giving specific status to domesticated forms whose genetic variation is minimal is a point of controversy, the detailed descriptions of domesticated species provided in Dorofeev et al. represent an important contribution. The full range of wheat diversity cannot be described by the abbreviated species concepts (6, 10, 11).

An English version of Dorofeev et al. will open a body of information to interested plant breeders, geneticists, genebank managers, botanists, archaeobotanists, and others working with wheat diversity. There are 1,242 infraspecific taxa that are catalogued with botanical descriptions, synonymy, taxonomic keys, geographic distribution, disease traits, origin, and history. As an example, the translation of the description of Triticum urartu is presented in Appendix 1. Over 3,000 names for the wheats are listed in the monograph’s index. While the original Russian-language publication provided only two pages of references, the translation also will include a full reference list with numerous publications not generally known outside of Russia.

 

5. FUNDING OF THE TRANSLATION PROJECT

Initial funding for DTP was provided by CIMMYT as a US$ 5,000 donation for fees associated with the Russian to English translation. The project is actively seeking other donations from public and private research institutions to fund the costs of scientific editing and publication (Morrison et al., 2000). The project fund is maintained at the Society for the Support of Research on Cultivated Plants in Gatersleben (GFK).

Interested donors can contribute as follows (banking instructions can be requested from H. Knüpffer, knupffer@ipk-gatersleben.de.):

  • Non-EEC citizens: by cheque payable to GFK-Gatersleben;
  • EEC citizens: by Eurocheque payable to GFK-Gatersleben;
  • German citizens: by Eurocheque payable to GFK-Gatersleben, or via bank transfer.

Donations should be addressed to:

Dorofeev Translation Project Fund, Mr. B. Eise

Gemeinschaft zur Förderung der Kulturpflanzenforschung e.V. (GFK)

Corrensstr. 3

D-06466 Gatersleben, Germany

 

6. PUBLICATION PLANS

The 323 text pages have been completely translated. Scientific editing by one of the monograph’s authors, Anna Filatenko, is currently underway. A revised reference list also is in process. Linguistic editing is scheduled to start in November 2001. Final editing and revisions will start in January 2002 and continue through the Fall of 2002. The projected publication date is set for early 2003.

The DTP plans to publish an affordable volume that will cost no more than $50. A publisher has not yet been chosen. Arrangements are planned to allow electronic publication of the taxonomic keys and other relevant portions of the translation on the GrainTax website.

The DTP has obtained permission from VIR to publish the translated version and to use photographs and figures from the original.

 

7. PROJECT VOLUNTEERS AND STAFF

The translation project was initiated in the summer of 1999 during the Percival Symposium "Wheat – Yesterday, Today and Tomorrow" at the University of Reading, UK (12). The committee overseeing the project includes the following individuals:

  • Helmut Knüpffer – Institute of Plant Genetics and Crop Plant Research (IPK), Gatersleben, Germany
  • Anna Filatenko (retired, VIR) – St. Petersburg, Russia
  • Laura Morrison – Oregon State University, Corvallis, Oregon, USA
  • Karl Hammer – Universität Gesamthochschule Kassel, Witzenhausen, Germany
  • Alexei Morgounov – CIMMYT, Almaty, Kazakhstan
  • Iva Faberová – Research Institute of Crop Production, Prague, Czech Republic

Irina Sokolova of the Komarov Botanical Institute (St. Petersburg, Russia) prepared the Russian to English translation of the text. Editorial responsibilities are allocated as follows: scientific editing by A. Filatenko; linguistic editing by Charles Jeffrey (St. Petersburg); final editing by A. Filatenko, K. Hammer, and L. Morrison; editorial supervision by H. Knüpffer.

 

ACKNOWLEDGEMENTS

The contribution of CIMMYT to the project fund is gratefully acknowledged. We also thank the "Gemeinschaft zur Förderung der Kulturpflanzenforschung" (GFK), Gatersleben, for hosting and maintaining the project fund. The work of A. Filatenko was carried out under the sponsorship of the Cooperative Agricultural Sciences Programme between Germany and the Russian Federation and funded by the German Ministry of Agriculture.

 

REFERENCES

[1] Dorofeev VF, Filatenko AA, Migushova EF, Udaczin RA, Jakubziner MM: Wheat. In: Dorofeev VF, Korovina ON (eds) Flora of Cultivated Plants, vol. 1. Leningrad, Russia, Kolos. 346 pp. (1979) (in Russian).

[2] Percival J: The Wheat Plant. Duckworth & Co, London, 463 pp. (1921).

[3] Bowden WM: The taxonomy and nomenclature of the wheats, barleys, and ryes and their wild relatives. Can. J. Bot. 37: 657–684 (1959).

[4] Schulz A: Die Geschichte der kultivierten Getreide. Halle, Verlag Louis Neberts (1913).

[5] Mac Key J: A plant breeder’s aspect on taxonomy of cultivated plants. Biol. Zentralbl. 107: 369-379 (1988).

[6] Kimber G, Sears ER: Evolution in the genus Triticum and the origin of cultivated wheat. In: Heyne EG (ed) Wheat and wheat improvement. 2nd ed. Madison, Amer. Soc. Agron., pp. 154–164 (1987).

[7] Morrison LA, Raup WJ: GrainTax synonymy table project: June 1999 progress report. Wheat Inform. Serv. 88: 52–56 (1999).

[8] Morrison LA, Raup WJ: GrainTax synonymy tables project: June 2000 Progress Report. Wheat Inform. Serv. 90: 54 (2000).

[9] Morrison LA, Faberová I, Filatenko AA, Hammer K, Knüpffer H, Morgounov A, Rajaram S: Call to support an English translation of the 1979 Russian taxonomic monograph of Triticum by Dorofeev et al. Wheat Inform. Serv. 90: 52–53 (2000).

[10] Morris R, Sears ER: The cytogenetics of wheat and its relatives. In: Quisenberry KS, Reitz LP (eds) Wheat and wheat improvement. Madison, Amer. Soc. Agron., pp. 19–87 (1967).

[11] Kimber G, Feldman M: Wild wheat: an introduction. Special Report 353. Columbia, Coll. Agric., Univ. Missouri (1987).

[12] Caligari PDS, Brandham PE (eds): Wheat taxonomy: the legacy of John Percival. The Linnean, Special Issue No. 3 (2001).

 

 

 

APPENDIX 1. T. URARTU Thum. EX GANDIL.

(extracted from the English translation of Dorofeev et al. (1979) (1))

Russian-to-English Translation: I. Sokolova, St. Petersburg

Editing: A. Filatenko, St. Petersburg; L. Morrison, Corvallis, Oregon, USA; H. Knüpffer, Gatersleben, Germany

 

T. urartu Thum. ex Gandil. 1972, Bot. Zhurn. 57, 2: 176; Tumanian 1938, Tr. Arm. Fil. AN SSSR, Ser. Biol. 2: 211, descr. ross. – T. armeniacum Thum. ex Flaksb. 1939, Opred. Nast. Khlebov: 27, nom. nud., non T. armeniacum Nevski 1933, non T. armeniacum Jakubz. 1938 et non T. armeniacum Makusch. ex Menabde 1945. – T. monococcum subsp. urartu (Thum.) A. et D. Löve 1961, Bot. Not. (Lund) 114, 1: 49, nom. illeg. – T. boeoticum Boiss. subsp. urartu (Thum.) Vav. 1964, Mirov. Resursy Khlebn. Zlakov: 21, nom. illeg. – T. boeoticum subsp. urartu (Thum.) V. Dorof. 1968, Euphytica, 17: 453, nom. illeg. – T. monococcum subsp. michaelii An. Fed. (Theod.) et Takht. ex Zhuk. 1968, Bot. Zhurn. 53, 4: 442, nom. nud. – T. michaelii Zhuk. 1971, Kult. Rast. Sorod.: 96, nom. nud.

Type: Armenian Republic, southeastern outskirts of Erevan near Vokhchaberd and Gekhadir villages, 30. VI. 1968, P. A. Gandilyan.

General characteristics. Seedlings violet or dark green. Plant prostrate at base. Culm thin, flexible, up to 145 cm high. Stem nodes violet. Leaves velvety pubescent or glabrous, linear-lanceolate (15 cm long, 0.7–1.0 cm wide). Leaf sheaths velvety pubescent or glabrous. Auricles white, ciliate. Spikes elongate (Fig. 3, a), narrow (7–9 cm long, 0.6–0.7 cm wide), fragile, spontaneously disarticulating. Spikelets two-awned, one-grained (less often two-grained). Lateral (two-rowed) side of spike approximately twice as wide as frontal side. Pubescence of rachis internode segments the same as for T. boeoticum. Glumes scabrous-tuberculate; more delicate than for T. boeoticum,. Keel tooth long with a broad base. Tooth of main lateral glume vein considerably less distinct than keel tooth and 8–10 times shorter than in T. boeoticum. Anthers short, 2.0–2.7 mm (Gandilyan, 1972).

T. urartu has a weaker stem than T. boeoticum: stem fracture strength at first internode is 456 g versus 2,000–3,000 g for T. boeoticum (Gradchaninova, 1967).

In Armenia, found on dry slopes of foothills.

All forms of T. urartu are winter forms. Heading stage is long when sown under irrigated conditions in Tashkent. Both in Armenia and in other native habitats in the Near and Middle East, T. urartu only occurs where T. boeoticum is present.

Compared to T. boeoticum, T. urartu shows less morning flowering. It also flowers in the evening (Zhukov, 1969). In Tashkent, T. urartu starts flowering before full emergence of the spike from the flag leaf (data of E. F. Migushova). In Armenia, its native land, T. urartu begins later than T. boeoticum (Araratyan and Surmenyan, 1939). T. urartu is separated from other diploid species by anther morphology as well (Dhaliwal and Johnson, 1976). Its anthers are unusually short (on the average 2.2mm, versus 3.6mm in T. boeoticum). They dehisce by a longitudinal cleft twisting into a spiral after flowering. Anthers of T. urartu are placed at the same level as the stigma; the lemma and palea open with difficulty (Araratyan and Surmenyan, 1939).

T. urartu, like other wild wheat species, stands out because of its high seed protein content (23.7- 25.0%) (Konarev et al., 1971). Lysine content is relatively high: 2.67-2.48% of total protein; 0.632-0.681mg per 100g of grain. On the average for three years, content of protein in grain of this species (reproduction in lowland Daghestan under irrigation) made up 19.1–23.7%, that of starch 53.0–58.0% (Jakubziner and Pokrovskaya, 1971).

Immunity. T. urartu is distinguished from T. boeoticum by its strong susceptibility to yellow (stripe) rust (Gulkanyan, 1938; Vavilov, 1957, 1964; Jakubziner, 1969). In the Tashkent region during 1968–1973, stripe-rust infections reached number 3 (on a 5-point scale) powdery mildew infections reached number 4 (Migushova, 1976). Sukhanberdina (1977) also found severe powdery mildew infections both in the seedling and adult plant stages.

Resistance to infection by races of brown and stem rust in the seedling stage proved to be stronger (type IV) than in T. boeoticum (Grigoryeva, 1975). Adult plants in field conditions were essentially resistant. M. M. Jakubziner (unpublished information) has noted that in his joint experiment with G. S. Turov in Southern Kazakhstan the degree of infection of adult plants by brown rust did not exceed 10%, although response to infection by the fungus was characterised as being of type IV.

Artificial infection by loose smut races, f. aestivi and f. duri, caused infection of 10.0 and 11.5% of spikes respectively (Yamaleev, 1974, Krivchenko et al., 1976). The fungal mycelium infected all the embryonic organs of the caryopsis. These researchers concluded that T. urartu was

distinctly different from T. boeoticum in having no embryo race-specific resistance to loose smut in all growth stages.

In general, T. urartu is not resistant to fungal diseases.

Geographical distribution. The species distribution range in Armenia is at southeastern outskirts of Erevan and around the environs of the villages Shorbulak, Gekhadir, Vokhchaberd, and Atsavan in the Abovyan district (after Gandilyan, 1976, refined).

In recent years, forms of T. urartu were reported from Iran (Jaaska, 1974), as well as from Turkey, Iraq and Lebanon (Johnson, 1975).

Origin. A number of researchers envision the phylogenetic development of wheat as forming a branched tree, which originated from a common ancestor. According to Flaksberger (1935), the ancestor is the progenitor for the diploid and polyploid wheat species. Nevski (1933) advanced the idea that the genus Triticum originated from the ancestral tribe Protohordeae, or more particularly from plants which had a paniculate inflorescence with 2-3 spikelets grouped at each rachis node. Sinskaya (1955) believed that the tribe Protohordeae consisted of perennial biotypes (2n = 14), which evolved and turned to annual growth habit, while gaining features of xerophily. Tzvelev (1976) concluded that the coasts of the ancient Tethys ("Ancient Mediterranean") were a region where highly specialised annuals evolved from festucoid grasses. These grasses then gave rise to the first diploid wheat. In our opinion, this ancestor could have been an ancient form of T. urartu.

Taxonomy. T. urartu was found by M. G. Tumanian in 1934 who named and published the species description in Russian in 1938. Its spikes were brought into the VIR collection by N. I. Vavilov, who identified them as T. armeniacum Thum. This name was published in 1939. The T. armeniacum epithet is illegitimate, since armeniacum was assigned to another species of two-grained wheats in 1933. In 1938 and 1945, the name T. armeniacum was used as a later homonym of T. araraticum. The epithet michaelii suggested by Fedorov and Takhtajan (1968) is also a later homonym of T. urartu. Thus, a certain confusion was introduced into nomenclature.

The diagnosis of T. urartu was published by Gandilyan (1972). For many years in the botanical and genetic literature, T. urartu has been regarded as a subspecies of T. boeoticum (Vavilov, 1964) or included in T. monococcum (Löve and Löve, 1961).

Key to the determination of varieties of T. urartu Thum. ex Gandil.

Glumes

Awns

Variety

Number of variety

white

white with black margins

red

red with black margins

black on background

same as glumes

black

white

red

+

–

–

–

–

–

+

–

spontaneoalbum

1

–

+

–

–

–

–

+

–

binartulutriru

2

–

–

+

–

–

–

+

–

spontaneorubrum

3

–

–

–

+

–

–

–

+

urartu

4

–

–

–

–

+

–

–

+

albinigricans

5

–

–

–

–

–

+

–

+

nigrum

6

Var. albinigricans Thum. ex Dorof. et A. Filat. 1979, in Dorof. et al., Kult. Fl. 1: 38. – T. urartu var. albonigricans Thum. 1938, Tr. Arm. Fil. AN SSSR, Ser. Biol. 2: 214, descr. ross. (5). A T. urartu var. spontaneoalbum glumis aristisque nigris differt.

Typus: Armenia. Reproductio stationis Daghestanskajae fulcracea, WIR, k-33871, 3. VI. 1968, leg. Soskov (WIR).

Geographical distribution. Armenia.

Var. binartulutriru Gandil. ex Dorof. et A. Filat. 1979, in Dorof. et al., Kult. Fl. 1: 38. – T. urartu var. binartulutriru Gandil. 1975, Tr. Arm. NIIZ, Ser. Pshenitsa (Echmiadzin), 3: 74, descr. ross. (2). Spica alba marginibus glumarum nigris.

Typus: Armenia, prope pagum Gegadir, 30. VI. 1968, leg. P. A. Gandiljan (WIR).

Geographical distribution. Armenia.

Var. nigrum Thum. ex Dorof. et A. Filat. 1979, in Dorof. et al., Kult. Fl. 1: 38. – T. urartu var. albonigricans Thum. 1938, l. c.: 214, descr. ross. (6). A T. urartu var. urartu glumis nigris differt.

Typus: Armenia. Reproductio stationis Daghestanskajae fulcracea, WIR, k-33870, 29. VI. 1968, leg. Soskov (WIR).

Geographical distribution. Armenia.

Var. spontaneoalbum Thum. ex Dorof. et A. Filat. 1979, in Dorof. et al., Kult. Fl. 1: 38. – T. urartu var. spontaneoalbum Thum. 1938, l. c.: 214, descr. ross. (1). Spica aristaque albae.

Typus: Armenia. 30. VI. 1968, leg. P. A. Gandiljan (WIR).

Geographical distribution. Armenia.

Var. spontaneorubrum Thum. ex Dorof. et A. Filat. 1979, in Dorof. et al., Kult. Fl. 1: 38. – T. urartu var. spontaneorubrum Thum. 1938, l. c.: 214, descr. ross. (3). Spica aristaque rubrae.

Typus: Armenia. Reproductio stationis Daghestanskajae fulcracea, WIR, k-33869, 3. VI. 1968, leg. Soskov (WIR).

Geographical distribution. Armenia.

Var. urartu. – T. urartu var. binarturutriru Gandil. 1972, Bot. Zhurn. 57, 2: 177. (4).

Geographical distribution. Armenia.