28. Акопян Ж.А. О некоторых биоморфологических особенностях прорастания Climacoptera crassa (Chenopodiaceaе) // Материалы межд. науч. конф. “Проблемы биоразнообразия Кавказа”. Махачкала. 2007. С. 48-49.
29. Akopian J.A., Gabrielyan I.G. and H. Freitag. Fossil fruits of Salsola L. s.l. and Halanthium K. Koch (Chenopodiaceae) from Lower Pleistocene lacustrine sediments in Armenia // Feddes Repertorium, Journal of Botany Taxonomy and Geobotany. 2008. Federal Republic of Germany. Band 119 (3-4). Р. 225-236.
30. Акопян Ж.А. Сохранение разнообразия флоры Армении в Ереванском ботаническом саду – история и перспективы // Материалы межд. конф. “Актуальные вопросы ботаники в Армении”. Ереван. 2008. С.56-61.
31. Акопян Ж.А. Обзор жизненных форм представителей семейства Chenopodiaceae Vent.
во флоре Армении // Материалы межд. конф. “Актуальные вопросы ботаники в Армении”.
Ереван. 2008. С. 56-61.
32. Акопян Ж.А., Айрапетян А.М. О некоторых особенностях морфологии соцветия, цветка и пыльцы Krasheninnikovia ceratoides (L.) Gueldenst. (Chenopodiaceae) // Материалы межд. науч. конф., посв. 100-летию П.И. Лапина “Проблемы современной дендрологии”.
Москва. 2009. С. 590-593.
33. Akopian J.A. Crop relative conservation on the plot “Flora and vegetation of Armenia of the Yerevan Botanical Garden”//Electronic J. Nat. Sci. 2009, 2 : 3-7. Available at http://www.epnet.com http://www.ejns.sci.am 34. Akopian J.A. Conservation of the aboriginal plant diversity at the plot “Flora and vegetation of Armenia” of the Yerevan Botanical Garden // “Plant conservation for the next decade – a cerebration of Kew’s 250th anniversary”. Abstracts. Royal Botanic Gardens. Kew. UK. 2009. P. 69.
35. Акопян Ж.А. Биолого-морфологическое исследование представителей рода Noaea (Chenopodiaceae) в Армении // Флора, растительность и растительные ресурсы Армении. 2009.
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39. Akopian J.A. On the biomorphology of Hablitzia tamnoides (Chenopodiaceae) // Proceedings of the Symposium. Caryophyllales: New Insights into the Phylogeny, Systematics and Morphological Evolution of the Order. Moscow. 2012. P. 37-40.
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Chenopodiaceae Chenopodiaceae Salsoloideae Chenopodioideae, Polycnemoideae : 29, 1-6, Caroxylon, Climacoptera, Halothamnus, Kali, Kaviria, Salsola s. str., Chenopodiaceae, Chenopodioideae Betoideae, (78,3%): Hablitzia tamnoides-, Microcnemum coralloides-, Halanthium :
Chenopodiaceae Biomorphological Peculiarities and Taxonomical Composition of the Chenopodiaceae Vent.
RESUMEThe Chenopodiaceae Vent. family is an important component of the flora and vegetation of South Transcaucasia (Armenia and Nakhichevan) where this family is represented by nearly 30% of the total genera composition, and 85.7% of genera and 80.3% of species composition of the Caucasus.
Nevertheless, a complete study of this family in ST region has not yet been carried out.
The purpose of our investigation is to identify and clarify the diversity of the Chenopodiaceae family of ST, using the methods of taxonomical, biomorphological and phytogeographical analysis.
The taxonomical revision of the family shows that it is represented in ST by 90 species and subspecies from 36 genera and 8 subfamilies. The subfamilies Salsoloideae and Chenopodioideae are the most numerous ones in the region, whereas the least numerous one is Polycnemoideae. The dominated genera include 1-6 species - 29, monotype genera are 5, with 13-14 species – only 2.
The check-list of the family based on our field investigations and herbarium analysis was made.
The data on the species’ altitudinal and geographical distribution, habitats, flowering and fructification periods, main synonyms as well the citation of type specimens are given. The presence of 15 species, not listed in the “Flora of Armenia” (1956) but reported from the area by the earlier authors, is confirmed, and 7 species, viz. Chenopodium novopokrovskianum (Aellen) Uotila, Camphorosma monspeliaca L., Caroxylon gemmascens (Pall.) Tzvelev, Salicornia perennans Willd., Suaeda salsa (L.) Pall., S. gracilis Moq. and S. ekimii Freitag & Adigzel ined., are newly recorded from Armenia.
Some species are restored from the synonyms or are transferred into them, 2 species (Salicornia europaea L., Gamanthus pilosus (Pall.) Bunge) are excluded, 3 new combinations are given, new locations of 32 species are found. Genera Caroxylon Thunb., Climacoptera Botsch., Halothamnus Botsch., Kali Mill., Kaviria Akhani et E.H. Roalson, Salsola s. str. previously included in genus Salsola s.l. are accepted for ST. The biomorphological characteristics of these genera, proving their independence, are provided. The keys to genera and species determination are given. The maps of all ST Chenopodiaceae species are presented.
The main taxonomic and biomorphic diversity of the Chenopodiaceae family is observed in the arid regions of ST, in the area of Armeno-Iranian floristic province, in Yerevan floristic region and in Nakhichevan as well as in Meghri and Darelegiss floristic regions of Armenia. In the northern part of ST, related to the Caucasian floristic province, has been observed a taxonomical diversity of subfamilies Chenopodioideae and Betoideae represented by perennials, annuals and some taxa of segetal and ruderal plants. Here, the richest one in Chenopodiaceae representatives is Sevan floristic region.
Since Yerevan is located in the center of a number of Chenopodiaceae species natural areas of the studied region, an inventory of the family in the city has been carried out. There has been recorded a high percentage of Chenopods participation in the urbanoflora, 15 genera and 35 species have been recorded, making up to 46.8% of the generic composition and 46.6% of the total number of the species in Yerevan floristic region of Armenia.
In the result of the chorological analysis of Chenopodiaceae family, 21 geographical elements and 10 enlarged chorotype groups have been determined. Most of ST Chenopodiaceae species (78.3%) are of Ancient Mediterranean origin. The prevalence of East Ancient Mediterranean (21.1%), Vorden Asian (17.7%), Irano-Turanian (16.7%) and Euro - Ancient Mediterranean (14.4%) elements is revealed. The level of Chenopodiaceae species endemism in ST is low (3.4%). Caucasian species Hablitzia tamnoides belongs to the relic of Tertiary subtropical flora, Minor Asian Microcnemum coralloides ssp. anatolicum to the relic of early Miocene, and species of genus Halanthium belong to the relic of the early Pleistocene. On the bases of the paleopalinological and relict data analysis it was assumed that the origin of the halophyte flora and vegetation and Chenopodiaceae distribution in the area of ST took place earlier (Eocene-Miocene) than the Quaternary period (Takhtajan, 1941, 1946;
The bimorphological and anthecological investigation of Chenopodiaceae representatives was carried out in nature and in culture in the Yerevan Botanic Garden. Life forms of Chenopodiaceae family were studied, the classification represented by 4 types and 12 groups of life forms was worked out. Chenopodiaceae in ST region is represented by the following biomorphotypes: phanerophytes;
xeromorphic orthotropic and plagiotropic chamaeophytes and nanochamaeophytes; clump- or cushionforming chamaeophytes and nanochamaeophytes with rooting plagiotropic shoots; caudex-forming chamaeophytes and nanochamaeophytes; hemicryptophytes with annual monocyclic monocarpic shoots and special type of storage roots; vine-like hemicryptophytes with scrambling annual, monopodial, extremely elongated shoots and fleshy storage roots; therophytes (including biennials with basal rosettes). Therophytes (59 species) and chamaeophytes (25 species) are the most numerous in ST, while hemicryptophytes - 5 species, phanerophytes - 1 species.
The analysis of the spectrum of sexual types and forms of Chenopodiaceae and their distribution by taxa, a trend towards the development of hermaphrodite (Betoideae, Salicornioideae, Salsoloideae), monoeceous (Chenopodioideae) and gynomonoeoceous (Suaedoideae) plants has been reveal. The following types of pollination have been studied: xenogamy, geytonogamy, autogamy, cleistogamy.
Anemophyly as the main way of the cross pollination has been observed in 70 species, entomophyly in 20 species. Most typical is a combination of two or more types and ways of pollination. Protogeny as a type of flower dichogamy in the studied area has been observed in 45 species of 18 genera and protandry - in 23 species of 13 genera.
Hypothetical ancestral biomorphs for Chenopodioideae/Betoideae accepted axyle tropical plants with open buds, with seeds without rest period and for Salicornioideae/Suaedoideae - halosucculente herbaceous plants – helophytes of periodically flooded low littorals and inland ancient lagoons of Tethyan zone. During adaptation to the seasonal climate, in ancestral biomorphes were differentiated zones of regeneration, the origin of which is associated with plant hypocopilar-epicotylar zone, corresponding to modern Betoideae roots and to semi-woody Camphorosmoideae/Salsoloideae caudex with buds at the soil level. Further evolution of Chenopodiaceae biomorphs in arid conditions went on towards spatial spreading of the regeneration zone in the vertical direction due to lignification of shoots, development of metameric (modular) structure and lifting buds over the soil surface, and in the horizontal direction due to development of rooting plant forms.
In the result of ecological and ecomorphological analysis of ST Chenopods, the following plant groups have been identified: hyperhalophytes, euhalophytes, hygrohalophites, haloglykophytes, glycophytes; argilophytes, psammophytes, petrophytes; xerophytes, xeromesophytes, mesophytes.
Most species (56, 62.2%) according to the altitudinal zonation are native to the deserts/semideserts, foothills and lower mountain zone, at the height of (450) 600-1200 m a.s.l., and 34 species (37.8%) occur up to 2200 m a.s.l. Within the spectrum of the ST Chenopodiaceae, species with high halotolerance amount to 19 species (21%), weed-ruderal and segetal - 36 species (40%). The group of environmentally labile (plastical) species consists of 56 species (62.2%) and only 32 species (37.8%) have a narrow ecological specialization. Ecological analysis shows a high degree of adaptability of the majority of representatives of ST Chenopodiaceae.
The endangered species of Chenopodiaceae have been analyzed according to the IUCN criteria, 16 species are included in the Red Book of the Republic of Armenia (2010). In the result of Chenopods investigation of the Specially Protected Natural Areas of Armenia, existence of threatened species in these areas was specified.
Chenopodiaceae 43 species of 28 genera have been tested for cultivation on the "Flora and Vegetation of Armenia" plot in Yerevan Botanic Garden. Of these 7 threatened species, viz. Kaviria cana ssp. cana, K. tomentosa ssp. tomentosa, Beta lomatogona, Bienertia cycloptera, Halanthium kulpianum, Kali tamamschjanae, Spinacia tetrandra have high indicators of acclimatization and reproduction in culture and can be recommended for a future reintroduction into the wild.
The groups of ST Chenopodiaceae valuable plants (medicinal, fodder, ornamental, food plants, crop wild relatives) are determined, some species are recommended for use for landscaping of the region and for testing as food plants.