CC BY
10
Turczaninowia 23, 4: 50-64 (2020) DOI: 10.14258/turczaninowia.23.4.5 http://turczaninowia.asu.ru
ISSN 1560-7259 (print edition)
TURCZANINOWIA
ISSN 1560-7267 (online edition)
Y^K 582.669.24
Is Holosteum glutinosum (M. Bieb.) Fisch. et C. A. Mey. (Caryophyllaceae: Alsinoideae) just a subtaxon of H. umbellatum L. or a distinct species?
A. V. Fateryga1*, V. V. Fateryga1, I. V. Sokolova2, S. A. Svirin3, A. V. Yena4, P. E. Yevseyenkov5
1 T. I. Vyazemsky Karadag Scientific Station - Nature Reserve of RAS - Branch of A. O. Kovalevsky Institute of Biology of the Southern Seas of RAS, Nauki str., 24, Kurortnoye, Feodosiya, Republic of Crimea, 298188, Russian Federation. E-mails: fater_84@list.ru*, valentina_vt@mail.ru
2 V. L. Komarov Botanical Institute of the Russian Academy of Sciences, Prof. Popova str., 2, St. Petersburg, 197376, Russian Federation. E-mail: isokolova@binran.ru
3 Urban Development Institute of the Sevastopol State University, Kornilova emb., 1, Sevastopol, 299011, Russian Federation. E-mail: sapsan7@mail.ru
4Academy of Bioresources and Environmental Management of V. I. Vernadsky Crimean Federal University, Agrarnoye, Simferopol, Republic of Crimea, 295492, Russian Federation. E-mail: an.yena@gmail.com
5 Independent researcher, Oktyabrskoy Revolyutsii pr., 32, kv. 496, Sevastopol, 299038, Russian Federation.
E-mail: fhunt@flora.crimea.ru
Keywords, distribution, lectotype, nomenclature, systematics, taxonomy, typification.
Summary. The genus Holosteum L. distributed mostly in temperate Eurasia has very confused taxonomy. Modern estimation of the species richness varies from one species worldwide to seven species occurring just in Eastern Europe. One of the most problematic species of the genus is H. glutinosum (M. Bieb.) Fisch. et C. A. Mey. usually treated as a subspecies or a variety of H. umbellatum L. On the base of extensive study of the protologues, type material, a rich variety of other herbarium specimens, photos of all relevant taxa, and living plants in the field we recognize H. glutinosum as a distinct species. This species has three characters unique in the genus Holosteum. pure green color of leaves and sepals, completely herbaceous bracts, and lanceolate leaves; additional diagnostic characters are 10 stamens, dense glandular pilosity of leaves, sepals, bracts, stem, and pedicels, and entire petals longer than sepals. The species also has statistically significant differences from H. umbellatum (longer leaves, longer and wider petals, and longer capsule) while occurring in sympatry with it. Two subspecies are recognized within H. glutinosum. H. glutinosum subsp. glutinosum distributed in Caspian Depression, Transcaucasia, Western and Middle Asia, and H. glutinosum subsp. liniflorum Fateryga, nom. et stat. nov. (= H. liniflorum Steven ex Fisch. et C. A. Mey. 1840 [non 1837], nom. illeg.), presumably endemic to the Crimea. The lectotypes are designated for the latter name and also for H. glandulosum Bertol., H. umbellatum var. pleiandrum Fenzl, H. imberbe J. Gay, nom. illeg. superfl., H. imberbe var. brachypetalum J. Gay, and H. imberbe var. macropetalum J. Gay. Reports of H. glutinosum from most European countries (other than Russia) are considered to be erroneous due to numerous misidentifications in herbaria and incorrect synonymy (e. g., with H. heuffelii Wierzb., which is actually conspecific with H. umbellatum). A key to the species and subspecies is provided; a total of six taxa are recognized in the genus. H. breistrofferi Greuter et Charpin, H. glutinosum subsp. glutinosum, H. glutinosum subsp. liniflorum, H. marginatum C. A. Mey., H. tenerrimum Boiss., and H. umbellatum.
Поступило в редакцию 09.07.2020 Submitted 09.07.2020
Принято к публикации 02.11.2020 Accepted 02.11.2020
*Corresponding author
Holosteum glutinosum (M. Bieb.) Fisch. et C. A. Mey. (Caryophyllaceae: Alsinoideae) - лишь субтаксон H. umbellatum L. или самостоятельный вид?
А. В. Фатерыга1, В. В. Фатерыга1, И. В. Соколова2, С. А. Свирин3, А. В. Ена4, П. Е. Евсеенков5
1 Карадагская научная станция им. Т. И. Вяземского - природный заповедник РаАН - филиал Института биологии южных морей им. А. О. Ковалевского РаАН, ул. Науки, 24, пгт Курортное, г. Феодосия, Республика Крым, 298188, Россия
2Ботанический институт им. В. Л. Комарова РаАН, ул. Проф. Попова, 2, г. Санкт-Петербург, 197376, Россия
3 Институт развития города Севастопольского государственного университета, наб. Корнилова, 1,
г. Севастополь, 299011, Россия
4 Академия биоресурсов и природопользования Крымского федерального университета им. В. И. Вернадского, пгт Аграрное, г. Симферополь, Республика Крым, 295492, Россия
5Независимый исследователь, пр. Октябрьской Революции, 32, кв. 496, г. Севастополь, 299038, Россия
Ключевые слова: лектотип, номенклатура, распространение, систематика, таксономия, типификация.
Аннотация. Род Holosteum L., распространенный преимущественно в умеренной зоне Евразии, характеризуется крайне запутанной таксономией. Современные оценки его видового богатства варьируют от всего одного вида в мире до семи видов только лишь в Восточной Европе. Одним из наиболее проблемных видов является H. glutinosum (M. Bieb.) Fisch. et C. A. Mey., который обычно рассматривают как подвид или разновидность H. umbellatum L. На основе изучения протологов, типовых образцов, большого количества других гербарных материалов, фотографий всех релевантных таксонов и живых растений в природе установлено, что H. glutinosum является самостоятельным видом. Данный вид обладает тремя признаками, уникальными в роде Holosteum: чисто-зеленый цвет листьев и чашелистиков, полностью травянистые листочки обертки соцветия и ланцетные листья. Дополнительными диагностическими признаками являются 10 тычинок, густое железистое опушение листьев, чашелистиков, листочков обертки соцветия, стебля и цветоножек, а также цельные лепестки, превышающие по длине чашелистики. Вид обладает статистически значимыми отличиями от H. umbellatum (более длинные листья, более длинные и широкие лепестки и более длинная коробочка) при совместном произрастании с последним. В составе H. glutinosum выделено два подвида: H. glutinosum subsp. glutinosum, распространенный в Прикаспийской низменности, Закавказье, Западной и Средней Азии, и H. glutinosum subsp. liniflorum Fateryga, nom. et stat. nov. (= H. liniflorum Steven ex Fisch. et C. A. Mey. 1840 [non 1837], nom. illeg.), предположительно являющийся эндемиком Крыма. Для последнего названия, а также для H. glandulosum Bertol., H. umbellatum var. pleiandrum Fenzl, H. imberbe J. Gay, nom. illeg. superfl., H. imberbe var. brachypetalum J. Gay и H. imberbe var. macropetalum J. Gay обозначены лектотипы. Указания H. glutinosum для стран Европы (кроме России) предположительно являются ошибочными и основанными на многочисленных неверных определениях материала в гербариях и ошибочной синонимии (например, с H. heuffelii Wierzb., который в действительности конспецифичен H. umbellatum). Приводится ключ для определения видов и подвидов рода; в общей сложности признается шесть таксонов: H. breistrofferi Greuter et Charpin, H. glutinosum subsp. glutinosum, H. glutinosum subsp. liniflorum, H. marginatum C. A. Mey., H. tenerrimum Boiss. и H. umbellatum.
Introduction
Holosteum L. (Caryophyllaceae: Alsinoideae) is a small genus of annual herbs native to temperate Eurasia (Hernandez-Ledesma et al., 2015) and North Africa (Greuter et al., 1984) with the type species H. umbellatum L. that has additionally naturalized in the United States of America, Canada, Argentina, and the Republic of South Africa (Rabeler, Hartman, 2005). Although the genus is not species-rich, its taxonomy is very confused and requires a revision
(Fateryga et al., 2017). Modern estimations on the number of species in the genus vary. World Flora Online (2020) accepts the one species H. umbellatum with four subspecies: H. umbellatum subsp. glutinosum (M. Bieb.) Nyman, H. umbellatum subsp. hirsutum (Mutel) Breistr., H. umbellatum subsp. viscosissimum (Celak.) Dvorak, and the type subspecies, while many other names in the genus Holosteum remain unresolved in this database. K. Marhold (2011) in Euro+Med PlantBase lists four species occurring in Europe and the Mediterranean
countries. Among them, H. umbellatum has five subspecies: those accepted in World Flora Online plus H. umbellatum subsp. tenerrimum (Boiss.) Greuter et Burdet. Three other species in Euro+Med PlantBase are H. marginatum C. A. Mey., which is a well recognized species (Fateryga et al., 2017), H. sperguloides Lehm., which actually belongs to the genus Spergularia (Pers.) J. Presl et C. Presl (World Flora Online, 2020), and H. subglutinosum Klokov.
P. Hernandez-Ledesma et al. (2015) reported the genus Holosteum to include three to four species worldwide without listing their names. A. V. Fateryga et al. (2017) supposed that there were four species: H. glutinosum (M. Bieb.) Fisch. et C. A. Mey., H. tenerrimum Boiss., H. umbellatum, and H. marginatum. One of the most "splitting" estimations was provided by N. N. Tzvelev (2012) who recognized six species within the territory of Eastern Europe. Among them, there were H. marginatum, H. glutinosum, H. umbellatum, H. klopotovii (Tzvelev) Tzvelev, H. syvaschicum Kleopow, and H. subglutinosum; in our opinion, however, the latter three represent different pilosity variations of H. umbellatum and do not merit the rank of species or even subspecies.
Holosteum glutinosum is the most problematic taxon in the genus. It is usually treated as a subspecies of H. umbellatum (Atlas Florae ..., 1983; Walters, Akeroyd, 1993; Marhold, 2011; World Flora Online, 2020) or even as a variety H. umbellatum var. glutinosum (M. Bieb.) J. Gay (Coode, 1967; Zernov, Sokolov, 2004), although the latter name is superfluous (see below). On the other hand, some modern Russian authors (Lazkov, 2006, 2012; Tzvelev, 2012) continue to treat H. glutinosum as a full species as it was traditionally accepted in Soviet literature (Muravyeva, 1936; Grossheim, 1945; Czerepanov, 1995). It is generally considered that H. (umbellatum subsp./var.) glutinosum has 10 stamens and glandular pubescence developed on the whole green parts of the plant, while H. umbellatum (s. str.) has 3-5 stamens and bare bracts and/or lower parts of the stem/pedicels (Muravyeva, 1936; Grossheim, 1945; Schischkin, 1947; Coode, 1967; Walters, Akeroyd, 1993). Some authors also add that the first taxon has petals 1.5-2 times longer than sepals (Muravyeva, 1936; Schischkin, 1947; Lazkov, 2006) and capsule longer than 6.5 mm (Coode, 1967) while the second one has petals a little longer than sepals and capsule not longer than 6 mm.
A. S. Zernov and I. V. Sokolov (2004) considered that all diagnostic characters mentioned for both
H. glutinosum and H. umbellatum varied within their range from northwest to southeast showing a cline and making them to reduce H. glutinosum to the variety of H. umbellatum. On the other hand, N. N. Tzvelev (2004) reported that H. glutinosum had herbaceous bracts and narrow leaves while H. umbellatum had membranous bracts and broader leaves and G. A. Lazkov (2006) reported that
H. glutinosum had entire petals while H. umbellatum had petals with dentate apical margin. These characters were overlooked by A. S. Zernov and
I. V. Sokolov (2004), as well as by earlier European authors (Coode, 1967; Walters, Akeroyd, 1993). Such a confusion with the diagnostic characters of these two taxa led us to an assumption that some of the authors erroneously assigned the most pubescent plants of H. umbellatum s. str. to H. (umbellatum subsp./var.) glutinosum and such a mistake could be partial reason for reducing the latter species to a subtaxon of the former one.
At the same time, we found in 2013 a peculiar population of H. glutinosum in the Crimea, which occurred in the same habitat with H. umbellatum and showed remarkably distinct characters from the latter. These plants had characters corresponding to H. liniflorum Steven ex Fisch. et C. A. Mey. 1840 [non 1837], nom. illeg. (Fischer et al., 1840; see also Steven, 1856), which is currently treated as a synonym ofH. umbellatum var. glutinosum (Zernov, Sokolov, 2004). Thus, the purpose of the present contribution is to ascertain the taxonomic status of H. glutinosum, as well as other taxa hitherto assigned to its synonyms, and to provide an identification key to all taxa known in the genus Holosteum.
Material and methods
Our study was based on traditional taxonomic methods. All protologues of the species of interest, as well as additional relevant references, were studied. The type material of most taxa and a large amount of additional specimens were examined in nine herbaria: CSAU, G (CHG, 2020), K (Kew Herbarium, 2020), KW, LE, MW (Seregin, 2020), P (MNHN, 2020), PHEO, and YALT, as well as in various digital herbaria via the Global Plants webservice (https://plants.jstor.org/). Photos of living plants were examined on the Plantarium web-site (Plantarium, 2007-2020) and on various other web-sites via the Google search (https://www. google.com/). We used the morphological method and examined the main important characters of Holosteum plants: leaf shape; leaf, bract, and sepal coloration; bract size and structure; petal size and
shape; capsule size; stem, pedicel, bract, and sepal pilosity. Distribution of the species was studied on the base of critically reviewed literature data, as well as the studied herbarium and photographic materials.
Living plants of H. glutinosum were studied in April 2016 in Zuya, Belogorsk District of the Republic of Crimea (45°03'39"N, 34°20'06"E), where they occurred in the same habitat with H. umbellatum. Thirty plants of each species were randomly selected and measured with a vernier caliper. The measured parameters were the shoot length, the upper leaf length and width, a petal length and width, and an opened capsule length. Then, the minimum, maximum, and mean values were calculated for each parameter. Confidence intervals of the mean values were calculated for 95 % confidence level (p = 0.05). Statistical significance of the differences between the mean values of two species was checked with Student's Z-test (Lakin, 1990). PCA analysis of the same plants was made with Statistica 7 software.
To ascertain what taxonomic rank is to be applied to H. glutinosum, we assumed that a subspecies is a population (or several populations) of the species with morphological differences recognized by taxonomists (Mayr, 1974). This assumption means that two subspecies cannot co-occur in the same place. At the same time, the assumption does not mean that any morphological difference between allopatric populations is the reason to recognize one of them as a separate subspecies (many such differences are unstable and associated with ecological conditions).
Lectotype designation for H. liniflorum Steven ex Fisch. et C. A. Mey. 1840 [non 1837] was performed by Ari Taponen (University of Helsinki, Finland).
Results and discussion
What is Holosteum glutinosum?
Studying of all available materials has revealed that H. glutinosum is a well recognized species clearly distinct from four other well recognized species in the genus (see the key below) by three unique characters. pure green color of leaves and sepals, completely herbaceous bracts, and lanceolate leaves. These differences, however, are generally overlooked by most authors except N. N. Tzvelev (2004). Four other species have leaves with glaucous tincture, bracts either with membranous margins or nearly/almost completely membranous, and ovate leaves. Additional diagnostic characters of H. glutinosum, which are
not unique, are 10 stamens, dense glandular pilosity of leaves, sepals, bracts, stem, and pedicels, and entire petals longer than sepals.
Holosteum glutinosum is quite variable across its geographical range (Figs 1-2). The plants from the Volga Region, close to the type locality of the species (Fig. 1A), and the Middle Asia (Fig. 1B-D) usually have rather erect stem and relatively small petals. Plants from Asia Minor (Fig. 1E) and Transcaucasia (Fig. 1F-G) are rather ascending while their petals are also not very large. The most remarkable plants can be found in the Crimea where they always have erect, sometimes very tall stem and peculiarly large petals (Fig. 2A-F). All diagnostic characters of H. glutinosum listed above, however, are constant across its range.
The species has statistically significant differences from H. umbellatum when occurring in the same habitat with it. These differences are longer leaves, longer and wider petals, and longer capsule (Table). Samples of the two species studied in the Crimea also form two well recognized clouds in PCA analysis (Fig. 3). Thus, H. glutinosum cannot be treated as a subspecies or a variety ofH. umbellatum. There is also a difference in phenology, which is well visible in Fig. 2B where these two species are shown together: H. glutinosum in flower and H. umbellatum in fruit. Another question, however, arises: is the name H. glutinosum correctly applied to the "green" species (i. e., the species with pure green color of leaves and sepals, completely herbaceous bracts, and lanceolate leaves)?
Holosteum glutinosum was originally described by F. A. Marschall von Bieberstein (1808: 344) as Arenaria glutinosa M. Bieb. The author did not specify the color of leaves and the structure of bracts (herbaceous or membranous) of his species. He, however, provided the data on the leaf shape: "foliis oblongis obtusiusculis" that corresponded rather to the "green" species. F. A. Marschall von Bieberstein (1808) also placed his description to the chapter "Decandria trigynia" that meant that the plant had 10 stamens, i. e., also corresponded to H. glutinosum in its current treatment (H. umbellatum has usually 5 stamens). Early authors, however, placed A. glutinosa to either synonyms or subtaxa of H. umbellatum ("glaucous" species). For example, E. Fenzl (1842) listed H. glutinosum among the synonyms of a glandularly pubescent variety H. umbellatum P [var.] pleiandrum Fenzl while obviously used another name H. liniflorum Steven ex Fisch. et C. A. Mey. for the "green" species (he, however, provided an incorrect reference to Fischer et al., 1838: 10, instead of Fischer et al., 1840: 52).
Fig. 1. Holosteum glutinosum (M. Bieb.) Fisch. et C. A. Mey. from various countries: A - Russia (Astrakhan Region); B - Kazakhstan (Jambyl Province); C - Uzbekistan (Tashkent); D - Kyrgyzstan (Chuy Province); E - Turkey (Igdir Province); F - Armenia (Yerevan); G - Azerbaijan (Nakhchivan Autonomous Republic). Photos by M. S. Knyazev (A), V. G. Kolbintsev (B), T. S. Tillaev (C), G. V. Chulanova (D and F), A. A. Lebedev (E), and N. K. Abbasov (G).
Fig. 2. Holosteum glutinosum (M. Bieb.) Fisch. et C. A. Mey. from the Crimea: A - habitat with flowering plants; B - flowering plant (on the left) together with fruiting plants of H. umbellatum L. (on the right); C - upper parts of flowering plants; D - inflorescence; E - part of a stem with two pairs of leaves; F - flower; G - opened capsule. Photos by P. E. Yevseyenkov (A), A. V. Fateryga (B and C), and S. A. Svirin (D-G).
Table
Morphometric parameters of the studied plants of Holosteum glutinosum (M. Bieb.) Fisch. et C. A. Mey. and H. umbellatum L. from the same habitat in the Crimea
Parameter n Holosteum glutinosum Holosteum umbellatum
min-max m ± M min-max m ± M
Shoot length, cm 30 8.8-30.0 15.7 ± 1.7 4.3-19.5 12.0 ± 1.6
*Upper leaf length, mm 30 6.0-37.5 14.3 ± 2.0 5.0-12.4 8.1 ± 0.7
Upper leaf width, mm 30 0.5-7.2 2.6 ± 0.5 1.2-7.0 3.3 ± 0.5
*Petal length, mm 15 6.5-8.4 7.3 ± 0.3 3.0-3.5 3.3 ± 0.1
*Petal width, mm 15 3.3-5.1 4.1 ± 0.3 0.8-1.4 1.2 ± 0.1
*Opened capsule length, mm 15 6.3-10.3 7.9 ± 0.5 5.0-8.1 5.9 ± 0.4
Note: n - sample size, min - minimum value, max - maximum value, m - mean value, M - confidence interval (p = 0.05); parameters with statistically significant differences of values (p < 0.01) are marked with an asterisk.
Fig. 3. PCA analysis for plants of Holosteum glutinosum (M. Bieb.) Fisch. et C. A. Mey. (A) and H. umbellatum L. (B) from the same habitat in the Crimea.
J. Gay (1845: 27), while publishing the diagnosis of H. umbellatum, indicated that he assigned all "glaucous" plants to this taxon ("H. glaucum, <...> bracteis parvis membranaceis, <...>") and treated H. glutinosum as a new variety H. umbellatum P [var.] glutinosum. The latter name, however, is nomenclaturally superfluous, since an earlier name at the same rank (H. umbellatum P [var.] pleiandrum) was cited as its synonym [Art. 52.1, 52.2(e), 52.4 of ICN (Turland et al., 2018)]. At the same time, J. Gay (1845: 37) created a new name for the "green" species ("H. virens, <...> bracteis parvis, herbaceis, <...>"), namely H. imberbe J. Gay, and included H. liniflorum in its synonyms. Both E. Fenzl (1842) and J. Gay (1845) stated that the number of stamens in H. umbellatum P [var.] pleiandrum/glutinosum was up to 10 ("6-10" and "7-10", respectively). Thus, their varieties may probably represent an unsatisfactory mixture of the
traits of various specimens (of both "glaucous" and "green" species).
Soviet authors (Muravyeva, 1936; Grossheim, 1945; Schischkin, 1947; Czerepanov, 1995) were different in their taxonomic point of view on these two taxa. They clearly separated H. glutinosum (plants with 10 stamens) from H. umbellatum (plants with 3-5 stamens) but only N. N. Tzvelev (2004) completed their diagnoses with bract structure and leaf shape. The lectotype of A. glutinosa was designated by G. A. Lazkov and A. S. Zernov (see Zernov, Sokolov, 2004). The plants of the lectotype (Fig. 4) definitely represent the "green" species and clearly correspond to both the original description (Marschall von Bieberstein, 1808) and the modern Soviet/Russian treatment of H. glutinosum, although A. S. Zernov and I. V. Sokolov (2004) themselves did not accept it as a full species (see above).
As it was stated in the Introduction, modern taxonomic databases such as Euro+Med PlantBase (Marhold, 2011) or World Flora Online (2020) treat H. glutinosum as a subspecies of H. umbellatum that is not congruent with their co-occurring in sympatry (see above). One more possible reason of such an incorrect treatment, besides previous incorrect treatments by the early authors, is commonly occurring misidentifications of H. umbellatum, or sometimes also H. tenerrimum or even H. marginatum, as H. glutinosum in most European herbaria (with the exception of some Russian ones such as MW, see Seregin, 2020). For example, the plants on only one of nine sheets stored as H. glutinosum in P (MNHN, 2020) are correctly identified, namely a gathering from Armenia (P04975719). Among eight other sheets, six ones (P04975720, P04975802, P05020761-P05020763, and P05439558) represent typical H. umbellatum collected by Russian botanist A. K. Becker in Sarepta (currently Volgograd, Russia), close to the
type locality of H. glutinosum, in the second half of the XIX century (this case again confirms that the name H. glutinosum was commonly misapplied to H. umbellatum by earlier researchers). One more sheet (P04975801) also represents H. umbellatum but collected in Serbia where H. glutinosum seems not to be present (see below). Another sheet from Armenia (P04975721) is actually H. marginatum.
The third possible reason for the incorrect treatment of H. glutinosum as a subtaxon of H. umbellatum is incorrect synonymy. Thus, Euro+Med PlantBase (Marhold, 2011) and World Flora Online (2020) report H. heuffelii Wierzb. (= H. umbellatum subsp. heuffelii (Wierzb.) Dostal) as a synonym of H. umbellatum subsp. glutinosum. This synonymy has probably come from the "Atlas Florae Europaeae" (1983) and it was also repeated in the "Flora Europaea" (Walters, Akeroyd, 1993). We studied the original material on H. heuffelii from Romania on the Global Plants web-service (HAL0118102 and HAL0118103) and can state without doubts that the plants on both sheets belong to H. umbellatum instead of H. glutinosum. Moreover, we failed to find any specimens of H. glutinosum from any European country except Russia (see also Greimler, 2001). Thus, the authors who treat H. glutinosum as a subtaxon of H. umbellatum may actually be not familiar with true H. glutinosum.
What is Holosteum liniflorum?
The name H. liniflorum Steven ex Fisch. et C. A. Mey. was originally published in F. E. L. Fischer et al. (1837). The authors applied this name to the "glaucous" species with 10 stamens occurred in the Crimea and the area adjacent to the Caspian Sea. Thus, H. liniflorum was listed there with a half-wrong description, which contained some characters of both "glaucous" and "green" species. Arenaria glutinosa was cited as a synonym of H. liniflorum (Fischer et al., 1837). Thus, the latter name is illegitimate according to the Art. 52.1, 52.2(e) of ICN (Turland et al., 2018).
Three years later the same authors published a new combination H. glutinosum applied to the "glaucous" species ("H. glaucum; <...> petalis oblongis subintegerrimis longitudine latitudineque sepala vix excedentibus") (Fischer et al., 1840: 52) instead of the previously published H. liniflorum. At the same time, they provided a new diagnosis for H. liniflorum Steven ex Fisch. et C. A. Mey. 1840 [non 1837]: "H. viride; <...> petalis obovatis integerrimis latitudine longitudineque sepala multo superantibus" (Fischer et al., 1840: 52). The latter
name was applied to the plants from the Crimea, which clearly belonged to the "green" species, although this name was again illegitimate according to the Art. 53.1 of ICN (Turland et al., 2018) due to the homonymy with H. liniflorum Steven ex Fisch. et C. A. Mey. 1837. Modern taxonomic databases such as Euro+Med PlantBase (Marhold, 2011) or World Flora Online (2020) usually provide only the "earlier" H. liniflorum (published in 1837) among the synonyms of H. umbellatum subsp. glutinosum but miss the name published in 1840. Only A. S. Zernov and I. V. Sokolov (2004) listed both the "earlier" and the "later" H. liniflorum, although they also did not take into account their different diagnoses and treated both names as synonyms of H. umbellatum var. glutinosum, nom. superfl.
We can confirm that the Crimean H. glutinosum (Fig. 2) has very peculiar large petals, resembling those of a flax ("liniflorum"), and merits its own name, however at the rank of subspecies. After extensive study of the literature, we are not aware that any legitimate name at subspecies rank has been previously published for the taxon that used to be named "H. liniflorum Steven ex Fisch. et C. A. Mey." 1840 [non 1837] and therefore a new name is proposed in the present paper (see below).
Taxonomy
Holosteum glutinosum (M. Bieb.) Fisch. et C. A. Mey. 1840, Index Sem. Hort. Petrop. 6: 52. = Arenaria glutinosa M. Bieb. 1808, Fl. Taur.-Caucas. 1: 344. = Holosteum liniflorum Steven ex Fisch. et C. A. Mey. 1837, Index Sem. Hort. Petrop. 3: 39, nom. illeg. superfl. [Art. 52.1, 52.2(e) of ICN (Turland et al., 2018)]. = Holosteum umbellatum ß [var.] pleiandrum Fenzl in Ledeb. 1842, Fl. Ross. (Ledeb.) 1: 3741. = Holosteum umbellatum ß [var.] glutinosum (M. Bieb.) J. Gay, 1845, Ann. Sci. Nat., Bot. ser. 3, 4: 33, nom. superfl. [Art. 52.1, 52.2(e), 52.4 of ICN (Turland et al., 2018)]. = Holosteum umbellatum subsp. glutinosum (M. Bieb.) Nyman, 1878, Consp. Fl. Eur. 1: 112.
Type locality: "in sabuletis deserti inter Astrachan et Kisljar" [Russia]. Lectotypus (Lazkov, Zer-
1 Lectotypus (Fateryga, hic designatus): "Ex sabulis deserti Cumani. a. 1798" (LE: LE01017379). Since E. Fenzl (1842) cited "H. an glutinosi var.? <...> C. Koch in Lin-naea XV, p. 708. Nr. 180" (Koch, 1841: 708) among synonyms of his H. umbellatum var. pleiandrum, the latter name in not a replacement name, and is not automatically typified [Art. 6.11-6.13 of ICN (Turland et al., 2018)]. It is typified here by the earlier designated lectotype of Arenaria glutinosa (Zernov, Sokolov, 2004).
nov in Zernov, Sokolov, 2004, Novosti Sist. Vyssh. Rast. 36: 109): "Ex sabulis deserti Cumani. a. 1798" (LE: LE01017379) (Fig. 4).
= Holosteum glandulosum Bertol. 1842, Misc. Bot. (Bertol.) 1: 11. = Holosteum imberbe J. Gay, 1845, Ann. Sci. Nat., Bot. ser. 3, 4: 37, nom. illeg. superfl. [Art. 52.1, 52.2(e) of ICN (Turland et al., 2018)]. = Holosteum imberbe a [var.] brachypeta-lum J. Gay, 1845, Ann. Sci. Nat., Bot. ser. 3, 4: 40.
Type locality: "Ex portu William in saxis con-glomeratis" [Iraq]. Lectotypus (Fateryga, hic designatus): "Colonel Chesney's Expedition to the Euphrates. No. 16. Port William, March 1836. Breccia rocks; rare" (left-hand specimen, BOLO) (Fig. 5).
Diagnosis: Leaves and sepals pure green, completely covered with dense glandular hairs as well as stem and pedicels; bracts green, completely herbaceous, glandularly pubescent; upper leaves lanceolate; stamens 10; petals entire, longer than sepals.
Notes: When publishing the name H. imberbe, J. Gay (1845) listed several other names of species rank as the synonyms of its varieties. Among them, H. glandulosum was cited as a synonym of H. imberbe a [var.] brachypetalum while H. lini-florum Steven ex Fisch. et C. A. Mey. 1840 [non 1837] was cited as a synonym of H. imberbe P [var.] macropetalum J. Gay. We are here typifying both H. imberbe and H. imberbe var. brachypetalum by the same element as H. glandulosum. The herbarium sheet stored in BOLO and inscribed by A. Bertoloni is no doubt the original material of H. glandulosum (Fig. 5). However, it consists of three plants of two different species: two left-hand plants belong to H. glutinosum while the right-hand one is H. um-bellatum. Therefore, one of the left-hand plants is designated as the lectotype (Fig. 5). To notice, the duplicates of the Colonel Chesney's Expedition to the Euphrates were distributed in different Herbaria, so the isolectotypes of H. glandulosum can be found elsewhere. For example, J. Gay (1845) reported that he had seen the F. R. Chesney's specimen no. 16 in the herbarium of J. P. B. Delessert, which is at present stored in G.
One more name, H. polygamum K. Koch, is dubious. It was treated as a synonym of H. umbellatum var. pleiandrum by E. Fenzl (1842), H. imberbe by J. Gay (1845), and H. glutinosum by O. A. Muravye-va (1936), although it is impossible to ascertain even whether it really belongs to the genus Holosteum due to its unclear diagnosis and the absence of the type material (see Zernov, Sokolov, 2004). T. A. Ady-lov (1971) even treated H. polygamum as an ac-
cepted species erroneously using this name instead of H. glutinosum while S. K. Czerepanov (1995) accepted both H. glutinosum and H. polygamum.
Holosteum glutinosum subsp. glutinosum
Diagnosis: Plants rather erect or often somewhat ascending; petals elliptic, usually about 1.5 times as long as sepals, usually not overlapping on margins.
Distribution: Caspian Depression, Transcaucasia, Western and Middle Asia. Records from European countries other than Russia seem to be erroneous and come from either incorrectly identified material or incorrect synonymization (e. g., with H. heuffelii, see above).
Holosteum glutinosum subsp. liniflorum
Fateryga, nom. et stat. nov. = Holosteum liniflorum Steven ex Fisch. et C. A. Mey. 1840, Index Sem. Hort. Petrop. 6: 52, nom. illeg. (homonym of Holosteum liniflorum Steven ex Fisch. et C. A. Mey. 1837) [Art. 53.1 of ICN (Turland et al., 2018)]. = Holosteum imberbe P [var.] macropetalum J. Gay, 1845, Ann. Sci. Nat., Bot. ser. 3, 4: 41.
Type locality: "Tauria" [Russia: Crimea]. Lecto-typus (Taponen, hic designatus): "Alifke ad Salgi-rum Taur. camp. in agris" (H: H1334769) (Fig. 6).
Diagnosis: Plants always erect; petals obovate, about 2 times as long as sepals, always distinctly overlapping on margins.
Distribution: Presumably endemic to the Crimea.
Note: Both H. liniflorum Steven ex Fisch. et C. A. Mey. 1840 [non 1837] and H. imberbe var. macropetalum are lectotypified here by the same specimen.
Key to species and subspecies of the genus Holosteum
1. Plants without glandular pubescence; stem and pedicels completely glabrous; leaves with or without simple hairs; bracts relatively large, about half as long as sepals or longer, herbaceous with membranous margins; stamens 10; plants of subalpine habitats and grassy mountain slopes at high elevation ....................................................... 2
+ Plants usually with glandular pubescence at least on lower part of stem or/and lower leaves; bracts usually very small, often unnoticeable, several times shorter than sepals, either completely herbaceous or membranous; stamens in various number; plants of various habitats, often ruderal or segetal ............. 3
Fig. 4. Lectotype of Arenaria glutinosa M. Bieb. (LE: LE01017379). Upper part of one plant is shown enlarged on the left top; arrow indicates a bract, which is completely herbaceous.
LE 01017379
2. Upper pair of leaves with pubescent margins; pedicels becoming pendant after flowering but then erect again when fruits mature; seeds usually longer
than 1 mm (Alps) ....................................................
.......................... H. breistrofferi Greuter et Charpin
+ Upper pair of leaves glabrous; pedicels always erect, even in early fruits; seeds usually about 1 mm
long (Crimea, Caucasus, Asia Minor to Iran) .........
.......................................H. marginatum C. A. Mey.
3. Leaves and sepals pure green, completely covered with dense glandular hairs as well as stem
and pedicels; bracts green, completely herbaceous, glandularly pubescent; upper leaves lanceolate;
stamens 10; petals entire, longer than sepals ....... 4
+ Leaves and sepals green with glaucous tincture; glandular pubescence varies in density, usually there are some areas without hairs either on stem/pedicels or leaves/sepals/bracts; bracts whitish, rather membranous, often glabrous; upper leaves ovate; stamens in various number; petals either entire or with dentate apical margin, of various length ...... 5
Fig. 5. Lectotype of Holosteum glandulosum Bertol. (left-hand specimen indicated by an arrow, BOLO). Courtesy of Alma Mater Studiorum University of Bologna - Sistema Museale di Ateneo - Orto botanico ed Erbario.
Fig. 6. Lectotype of Holosteum liniflorum Steven ex Fisch. et C. A. Mey. 1840 [non 1837], nom. illeg. (H: H1334769).
H1334769
4. Plants rather erect or often somewhat ascending; petals elliptic, usually about 1.5 times as long as sepals, usually not overlapping on margins (Caspian Depression, Transcaucasia, Western and Middle Asia).....H. glutinosum (M. Bieb.) Fisch. et
C. A. Mey. subsp. glutinosum
+ Plants always erect; petals obovate, about 2 times as long as sepals, always distinctly overlapping
on margins (Crimea) .............H. glutinosum subsp.
liniflorum Fateryga, nom. et stat. nov.
5. Stamens 10; petals entire, about 2 times as long as sepals, distinctly overlapping on margins (Asia Minor, also reported from Greece by Willing, Willing, 2009) ......................H. tenerrimum Boiss.
+ Stamens usually 5; petals usually with dentate apical margin, about as long as sepals or slightly longer, not overlapping on margins (most part of Europe, west of North Africa, Caucasus, Western and Middle Asia; introduced to North and South America and South Africa) ........H. umbellatum L.
Conclusions
The genus Holosteum comprises five well recognized species: H. breistrofferi, H. glutinosum, H. marginatum, H. tenerrimum, and H. umbellatum. Among them, H. glutinosum has unique characters such as pure green color of leaves and sepals, completely herbaceous bracts, and lanceolate leaves. The distribution of this species includes the Crimea, Caspian Depression, Transcaucasia,
Western and Middle Asia. Reports of the species from most European countries (other than Russia) are doubtful and therefore its detailed range requires further study. Two subspecies are recognized within H. glutinosum: H. glutinosum subsp. glutinosum distributed over the most part of the species range and H. glutinosum subsp. liniflorum presumably endemic to the Crimea.
Acknowledgments
Namig K. Abbasov (Nakhchivan, Azerbaijan), Galina V. Chulanova (Volkhov, Russia), Alexander A. Lebedev (Moscow, Russia), Mikhail S. Knyazev (Yekaterinburg, Russia), Vladimir G. Kolbintsev (Taraz, Kazakhstan), and Tulkin S. Tillaev (Tashkent, Uzbekistan) provided photos of H. glutinosum and a permission to publish them. Umberto Mossetti and Cristina Nisi (Bologna, Italy) provided a scan of the type material of H. glandulosum and the permission to publish it. Alexander N. Sennikov (Helsinki, Finland) made some valuable comments on the manuscript. Ari Taponen (Helsinki, Finland) selected a specimen for the lectotype designation for H. liniflorum.
The work of A. V. Fateryga and V. V. Fateryga was a part of the State research project No. AAAA-A19-119012490044-3 of the A. O. Kovalevsky Institute of Biology of the Southern Seas of RAS. The work of I. V. Sokolova was a part of the State research project No. AAAA-A19-119031290052-1 of the V. L. Komarov Botanical Institute of RAS.
REFERENCES
Adylov T. A. 1971. Holosteum L. In: Opredelitel rasteniy Sredney Azii [Key to plants of Middle Asia]. Vol. 2. Ed. S. S. Kovalevskaya. Tashkent: Fan. Pp. 238-239. [In Russian] (Адылов Т. А. Holosteum L. // Определитель растений Средней Азии. Т. 2. Под ред. С. С. Ковалевской. Ташкент: Фан, 1971. С. 238-239).
Atlas Florae Europaeae. Distribution of vascular plants in Europe. 1983. Vol. 6. Eds. J. Jalas, J. Suominen. Helsinki: The Committee for Mapping the Flora of Europe & Societas Biologica Fennica Vanamo. 176 pp.
CHG. 2020. Catalogue des herbiers de Genève (CHG). Genève: Conservatoire & Jardin botaniques de la Ville de Genève. URL: http://www.ville-ge.ch/musinfo/bd/cjb/chg (Accessed 29 April 2020).
Coode M. J. E. 1967. Holosteum L. In: Flora of Turkey and the East Aegean Islands. Vol. 2. Ed. P. H. Davis. Edinburgh: Edinburgh University Press. Pp. 85-87.
Czerepanov S. K. 1995. Vascular plants of Russia and adjacent states (the former USSR). Cambridge, New York & Melbourne: Cambridge University Press. x + 516 pp.
Fateryga A. V., Svirin S. A., Yevseyenkov P. E., Yena A. V. 2017. On the presence of Holosteum marginatum C. A. Mey. (Caryophyllaceae: Alsinoideae) in the Crimea. Turczaninowia 20, 2: 23-30. [In Russian] (Фатерыга А. В., Свирин С. А., Евсеенков П. Е., Ена А. В. О произрастании Holosteum marginatum C. A. Mey. (Caryophyllaceae: Alsinoideae) в Крыму // Turczaninowia, 2017. Т. 20, № 2. С. 23-30). DOI: 10.14258/turczaninowia.20.2.3
Fenzl E. 1842. Ordo XVIII. Alsineae Bartl. In: C. F. Ledebour. Flora rossica sive enumeratio plantarum in totius Imperii Rossici provinciis europaeis, asiaticis, et americanis hucusque observatarum. Vol. 1. Stuttgart: Librariae E. Schweizerbart. Pp. 337-420. DOI: 10.5962/bhl.title.6606
Fischer F. E. L., Meyer C. A., Avé-Lallemant J. L. E. 1840. Index sextus seminum, quae Hortus Botanicus Imperialis Petropolitanus pro mutua commutatione offert. Accedunt animadversiones botanicae nonnullae. St. Petersburg: Ex typis Academiae Caesareae Petropollitanae. 67 pp.
Fischer F. E. L., Meyer C. A., Trautvetter E. R. 1837. Index tertius seminum, quae Hortus Botanicus Imperialis Petropolitanus pro mutua commutatione offert. Accedunt animadversiones botanicae nonnullae. St. Petersburg: Ex typis Christiani Hintze. 48 pp.
Fischer F. E. L., Meyer C. A., Trautvetter E. R. 1838 ("1837"). Index quartus seminum, quae Hortus Botanicus Imperialis Petropolitanus pro mutua commutatione offert. Accedunt animadversiones botanicae nonnullae. St. Petersburg: Ex typis Academiae Caesareae Petropollitanae. 52 pp.
Gay J. 1845. Holostei, caryophyllearum alsinearum generis, monographia. Ann. Sci. Nat., Bot. sér. 3, 4: 23-44.
Greimler J. 2001. Holosteum umbellatum (Caryophyllaceae) in Österreich. Neilreichia 1: 57-70.
Greuter W., Burdet H. M., Long G. 1984. Med-Checklist. A critical inventory of vascular plants of the circum-mediterranean countries. Vol. 1. Geneve: Conservatoire et Jardin botaniques & Berlin: Secretariat Med-Checklist, Botanischer Garten und Botanisches Museum Berlin-Dahlem. xvi + 330 pp.
Grossheim A.A. 1945. Flora Kavkaza [Flora ofthe Caucasus], 2nd ed. Vol. 3. Baku: Academy of Sciences ofAzer-baijanian SSR Press. viii + 494 + xvii pp. [In Russian] (Гроссгейм А. А. Флора Кавказа. 2-е изд. Т. 3. Баку: Изд-во АН Азербайджанской ССР, 1945. viii + 494 + xvii с.).
Hernández-Ledesma P., Berendsohn W. G., Borsch T., von Mering S., Akhani H., Arias S., Castañeda-Noa I., Eggli U., Eriksson R., Flores-Olvera H., Fuentes-Bazán S., Kadereit G., Klak C., Korotkova N., Nyffeler R., Ocam-po G., Ochoterena H., Oxelman B., Rabeler R. K., Sanchez A., Schlumpberger B. O., Uotila P. 2015. A taxonomic backbone for the global synthesis of species diversity in the angiosperm order Caryophyllales. Willdenowia 45(3): 281-383. DOI: 10.3372/wi.45.45301
Kew Herbarium. 2020. The Herbarium Catalogue. Kew: Royal Botanic Gardens. URL: http://www.kew.org/ herbcat (Accessed 29 April 2020).
Koch C. 1841. Catalogus plantarum, quas in itinere per Caucasum, Georgiam, Armeniamque annis 1836 et 1837 collegit. Linnaea 15: 705-723.
Lakin G. F. 1990. Biometriya [Biometry]. Moscow: Vysshaya Shkola. 352 pp. [In Russian] (Лакин Г. Ф. Биометрия. М.: Высшая школа, 1990. 352 с.).
Lazkov G. A. 2006. Semeystvo gvozdichnyye (Caryophyllaceae) vo flore Kyrgyzstana [Family Caryophyllaceae in the flora of Kyrgyzstan]. Moscow: KMK Scientific Press Ltd. 272 pp. [In Russian] (Лазьков Г. А. Семейство гвоздичные (Caryophyllaceae) во флоре Кыргызстана. М.: Товарищество научных изданий КМК, 2006. 272 с.).
Lazkov G. A. 2012. Holosteum L. In: Caucasian flora conspectus. Vol. 3, Pt. 2. Ed. A. L. Takhtajan. St. Petersburg & Moscow: KMK Scientific Press Ltd. Pp. 163-164. [In Russian] (Лазьков Г. А. Holosteum L. // Конспект флоры Кавказа. Т. 3, ч. 2. Под ред. А. Л. Тахтаджяна. СПб.-М.: Товарищество научных изданий КМК, 2012. С. 163-164).
Marhold K. 2011. Caryophyllaceae. In: Euro+Med PlantBase - the information resource for Euro-Mediterranean plant diversity. Berlin: Botanic Garden and Botanical Museum Berlin-Dahlem. URL: http://ww2.bgbm.org/Euro-PlusMed/ (Accessed 29 April 2020).
Marschall von Bieberstein F. A. 1808. Flora taurico-caucasica exhibens stirpes phaenogamas, in Chersoneso Taurica et regionibus caucasicis sponte crescentes. Vol. 1. Kharkov: Typis Academicis. vi + 429 pp. DOI: 10.5962/ bhl.title.10825
Mayr E. 1974. Populations, species, and evolution. Moscow: Mir. 460 pp. [In Russian] (Майр Э. Популяции, виды и эволюция. М.: Мир, 1974. 460 с.).
MNHN. 2020. The vascular plants collection (P) at the Herbarium of the Muséum National d'Histoire Naturelle (MNHN- Paris). Version 69.166. Occurrence dataset. URL: https://doi.org/10.15468/nc6rxy (Accessed via GBIF.org 29 April 2020).
Muravyeva O. A. 1936. Holosteum L. In: Flora SSSR [Flora ofthe USSR]. Vol. 6. Ed. B. K. Schischkin. Moscow & Leningrad: Academy of Sciences of USSR Press. Pp. 466-471. [In Russian] (Муравьева О. А. Holosteum L. // Флора СССР. Т. 6. Под ред. Б. К. Шишкина. М.-Л.: Изд-во АН СССР, 1936. С. 466-471).
Plantarium: open on-line atlas and key to plants and lichens of Russia and neighbouring countries. 2007-2020. [In Russian] (Плантариум: открытый онлайн атлас-определитель растений и лишайников России и сопредельных стран, 2007-2020). URL: http://www.plantarium.ru/ (Accessed 29 April 2020).
Rabeler R. K., Hartman R. L. 2005. Holosteum. In: Flora of North America North of Mexico. Vol. 5. Ed. Flora of North America Editorial Committee. New York & Oxford: Oxford University Press. Pp. 94-95. URL: http://beta. floranorthamerica.org/Holosteum (Accessed 29 April 2020).
Schischkin B. K. 1947. Caryophyllaceae Juss. In: E. W. Wulff. Flora Kryma [Flora of the Crimea]. Vol. 2, Fasc. 1. Moscow & Leningrad: OGIZ & Selkhozgiz. Pp. 117-171. [In Russian] (Шишкин Б. К. Caryophyllaceae Juss. // Е. В. Вульф. Флора Крыма. Т. 2, вып. 1. М.-Л.: ОГИЗ-Сельхозгиз, 1947. С. 117-171).
Seregin A. P. (ed.). 2020. Moscow University Herbarium (MW). Version 1.129. Moscow: Lomonosov Moscow State University. Occurrence dataset. URL: https://doi.org/10.15468/cpnhcc (Accessed via GBIF.org 29 April 2020).
Steven Ch. von. 1856. Verzeichniss der auf der taurischen Halbinsel wildwachsenden Pflanzen. Bull. Soc. Imp. NaturalistesMoscou 29(2): 234-334.
Turland N. J., Wiersema J. H., Barrie F. R., Greuter W., Hawksworth D. L., Herendeen P. S., Knapp S., Kusber W.-H., Li D.-Z, Marhold K, May T. W., McNeill J., Monro A. M, Prado J., Price M. J., Smith G. F. (eds.). 2018.
International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum Vegetabile 159. Glashütten: Koeltz Botanical Books. DOI: 10.12705/Code.2018
Tzvelev N. N. 2004. Holosteum L. In: Flora Europae Orientalis [Flora of Eastern Europe]. Vol. 11. Ed. N. N. Tzvelev. Moscow & St. Petersburg: KMK Scientific Press Ltd. Pp. 171-173. [In Russian] (Цвелев Н. Н. Holosteum L. // Флора Восточной Европы. Т. 11. Под ред. Н. Н. Цвелева. М.-СПб.: Товарищество научных изданий КМК, 2004. С. 171-173).
Tzvelev N. N. 2012. Holosteum L. In: Conspectus florae Europae Orientalis [Conspectus of the flora of Eastern Europe]. Vol. 1. Ed. N. N. Tzvelev. St. Petersburg & Moscow: KMK Scientific Press Ltd. Pp. 215-216. [In Russian] (Цвелев Н. Н. Holosteum L. // Конспект флоры Восточной Европы. Т. 1. Под ред. Н. Н. Цвелева. СПб.-М.: Товарищество научных изданий КМК, 2012. С. 215-216).
Walters S. M., Akeroyd J. R. 1993. Holosteum L. In: Flora Europaea, 2nd ed. Vol. 1. Eds. T. G. Tutin, N. A. Burges, A. O. Chater, J. R. Edmondson, V. H. Heywood, D. M. Moore, D. H. Valentine, S. M. Walters, D. A. Webb. Cambridge: Cambridge University Press. P. 164.
Willing R., Willing E. 2009. Notizen Griechenland Nr. 3. URL: http://www.willing-botanik.de/Notizen%20GR/ pdf-notizenGR/Notizen%20Griechenland%203.pdf (Accessed 29 April 2020).
World Flora Online. 2020. URL: http://www.worldfloraonline.org/ (Accessed 5 June 2020). Zernov A. S., Sokolov I. V. 2004. Genus Holosteum L. (Caryophyllaceae) in flora Caucasi. Novosti sistematiki vysshikh rasteniy [Novit. Syst. Pl. Vasc.] 36: 106-111. [In Russian] (Зернов А. С., Соколов И. В. Род Holosteum L. (Caryophyllaceae) во флоре Кавказа // Новости сист. высш. раст., 2004. Т. 36. С. 106-111).
