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14Новости систематики высших растений 2020 JLJ ISSN 0568-5443 (print)
Novitates Systematicae Plantarum Vascularium 51: 13-17 | |ssn 2687-1564 (onLine)
Morphology, nomenclature and distribution of Bassia montícola (Chenopodiaceae-Amaranthaceae), a poorly known species from Western Asia
Морфология, номенклатура и распространение Bassia montícola (Chenopodiaceae-Amaranthaceae), малоизученного вида из Западной Азии
Alexander P. Sukhorukov1*, M. A. Kushunina2
1 Department of Higher Plants, Biological Faculty, Lomonosov Moscow State University
Leninskiye Gory, 1/12, Moscow, 119234, Russia [email protected]
2 Department of Plant Physiology, Biological Faculty, Lomonosov Moscow State University
Leninskiye Gory, 1/12, Moscow, 119234, Russia
*Corresponding author
https://doi.org/10.31111/novitates/2020.51.13
А. П. Сухоруков1*, М. А. Кушунина2
1 Московский государственный университет имени
М. В. Ломоносова, биологический факультет, кафедра высших растений
Ленинские горы, 1, стр. 12, Москва, 119234, Россия [email protected]
2 Московский государственный университет имени
М. В. Ломоносова, биологический факультет, кафедра физиологии растений
Ленинские горы, 1, стр. 12, Москва, 119234, Россия
*Автор для переписки
Abstract. Kochia monticola was previously considered as a synonym for the widely distributed Irano-Turanian Panderia pilosa. After the merger of Kochia and Panderia with Bassia based on molecular phylogeny, K. monticola remained a synonym of Bassia pilosa. We claim that Bassia monticola, a name proposed by Kuntze (1891) for K. monticola, should be separated from B. pilosa based on morphological characters and localised distribution in mountainous regions of Iran, Iraq, Syria, and Lebanon at altitudes 1800-2600 m a. s. l.
Keywords: Bassia, Western Asia, taxonomy, Camphorosmoideae, Chenopodiaceae-Amaranthaceae.
Аннотация. Ранее Kochia monticola считали синонимом широко распространенного ирано-туранского вида Panderia pilosa. После объединения родов Kochia и Panderia с родом Bassia на основе молекулярно-филогенетических данных K. monticola рассматривается соответственно как синоним Bassia pilosa. В настоящей работе мы показали, что вид Bassia monticola (название, предложенное Kuntze (1891) для K. monticola) должен рассматриваться отдельно от B. pilosa, поскольку характеризуется хорошими морфологическими отличиями и имеет узкое распространение в горных районах Ирана, Ирака, Сирии и Ливана на высотах 1800-2600 м над ур. м.
Ключевые слова: Bassia, Западная Азия, таксономия, Camphorosmoideae, Chenopodiaceae-Amaranthaceae.
The systematic position and taxonomy of the Camphorosmoideae Ulbr. has been re-evaluated after extended molecular studies, and Bassia All. has undergone the most drastic taxonomic changes (Kadereit, Freitag, 2011). In its current composition, it encompasses ~20 Eurasian and African species, including the members of commonly accepted genera Chenoleoides Botsch., Kirilowia Bunge, Kochia Schrad., Londesia Fisch. et C. A. Mey., and Panderia Fisch. et C. A. Mey. All native American and Australian members as well as some Eurasian species previously included in Bassia now belong to other genera (Cabrera et al., 2009, 2011; Kadereit, Freitag, 2011). The most significant characters delimiting Bassia from allied genera are leaf anatomy and, in
Поступила в редакцию | Submitted: 10.06.2020
some taxa, perianth morphology at the fruiting stage (Cabrera et al., 2009; Kadereit, Freitag, 2011; Freitag, Kadereit, 2014; Sukhorukov, 2014). Bassia species are frequently very much alike in morphology but different in other (anatomical or physiological) characters, which support their different position on the phylogenetic trees (Akhani, Khoshravesh, 2013).
In Western Asia (Armenia, Azerbaijan, Bahrain, Georgia, Iran, Iraq, Israel/Palestine, Jordan, Kuwait, Lebanon, Oman, Qatar, Saudi Arabia, Syria, Turkey, UAE, and Yemen), Bassia is represented by 11 native and one alien species (Akhani, Khoshravesh, 2013; Sukhorukov, Akopian, 2013; Sukhorukov et al., 2016). Of the 11 native taxa, 10 are present in Iran (Akhani, Kho-
Принята к публикации | Accepted: 07.12.2020
shravesh, 2013; Sukhorukov, unpublished data), and the number of species continuously decreases northwards (Sukhorukov, Akopian, 2013) and southwards (Sukhorukov et al., 2016; Taifour, El-Oqlah, 2017). Bassia pilosa (Fisch. et C. A. Mey.) Freitag et G. Kadereit is easily distinguished from other species in the region by the vertical embryo position, a character previously considered as genus-specific in Panderia (e. g., Iljin, 1936; Hedge, 1997). Sukhorukov et al. (2016) stated that Bassia pilosa described from Talysh (eastern Azerbaijan) is clearly heterogeneous in the area and presented arguments for the acceptance of B. monticola (Boiss.) Kuntze in a specific rank proposed by Kuntze (1891). The latter species was originally described as Kochia monticola Boiss. (Boissier, 1846) and has sometimes been considered as a synonym for Panderia pilosa Fisch. et C. A. Mey. (= Bassia pilosa) (Aellen, 1967; Hedge, 1997, 2001). Other authors accepted Bassia monticola (usually as Panderia monticola (Boiss.) Bunge or Kochia monticola) as a separate species (e. g., Post, 1932; Ul-brich, 1934; Mouterde, 1966; Scott, 1978) or did not mention it in accounts (Zohary, 1966; Heller, Heyn, 1994; Haber, Semaan, 2007).
Due to misidentification of Bassia pilosa and B. monticola, it is difficult to discern which records actually belong to B. pilosa s. str. or to B. monticola in the areas where both species may be present. To date, the precise distribution of B. monticola is still unknown since the species is very poorly represented in the European herbaria. While preparing a treatment of Chenopodiaceae for the "Flora of Iraq" project, a large number of Bassia specimens from Western Asia were revised, which provided deeper insight into the morphology and choro-logy of B. monticola.
Material and methods
Herbarium specimens were examined and revised (if necessary) in the herbaria B, BM, BR, E, FI, G, H, HUJ, K, LE, M, MSB, MW, MHA, P, and W. The map was prepared using the online tool simplemappr.net (Shorthouse, 2010). Prior to scanning electron microscopy (SEM), perianths with soft tissues (hairs) were dehydrated in aqueous ethyl alcohol solutions of increasing concentrations and then in alcohol-acetone solutions and pure acetone according to a described procedure (Sukhorukov, Zhang, 2013; Sukhorukov, 2014). SEM observations were made with a JSM-6380 microscope (JEOL Ltd., Japan) at 15 kV after critical-point drying and sputter coating with gold-palladium.
Results and Discussion
Bassia monticola (Boiss.) Kuntze, 1891, Revis. Gen. Pl. 2: 547. = Kochia monticola Boiss. 1846, Diagn.
Pl. Orient. Nov. 1 (7): 82; Moq. 1849, in DC., Prodr. 13 (2): 133. = Panderia monticola (Boiss.) Bunge, 1880, Mem. Acad. Imp. Sci. St.-Petersbourg, ser. 7, 27, 8: 9 (Pfl.-Geogr. Beitracht. Fam. Chenop.). — Lec-totype (Sukhorukov, 2014: 349): Persia australis [Iran, Kohgiluyeh and Boyer Ahmad Prov.], in locis alpis Kuh-Daena [Kuh-e Dena], ubi nomades tentoria sua constituent, 21 VII 1842, Th. Kotschy, № 521.715 (G!; isolectotypes — BM001190958!; E00296928!; FI011279!; HAL0107073 — photo!; JE00026166!; K0000898880! [upper half of the sheet]; LE00011719!; LE00011720!; MO; P04907854! [upper right specimens]; WAG0004061!).
- Pentodon barbatus Ehrenb. 1879, in Boiss., Fl. Orient. 4 (2): 925, nom. inval., in syn.
Annual pubescent herb up to 15(20-25) cm, basally branched; stems ascendant or prostrate, whitish or reddish, glabrescent; leaves alternate, loosely arranged (except those in the lower part of the stem), filiform to lanceolate (usually linear), 10-15 x 1-2(2.5) mm, obtuse to acutish, with brachyblasts in their axils; inflorescence dense, bracts exceeding the flower clusters, up to 1 cm long (in the base of the inflorescence), continuously diminishing in size towards the inflorescence apex; flowers 1-3 in leaf axils, sessile; perianth cylindrical, 2-3 mm long in flowering stage, increasing up to 5 mm in fruiting; its base with tufts of simple hairs; perianth tube white, slightly pubescent or pilose and glabres-cent, sometimes almost glabrous; teeth 5, hairy at the top, with green dots at the flowering stage and tubercles at the fruiting stage (wing-like outgrowths absent); stamens 5, protruding, anthers 1.1-1.25 mm long; fruit compressed, ~1.5 mm long with easily rupturing white pericarp; seed with vertical horseshoe-shaped embryo and abundant perisperm.
Note. Boissier (1879) reported a horizontal embryo position in younger fruits. This statement is erroneous, perhaps due to the fact that almost all material available at that time was collected in blooming or early fruiting stages when the seed embryo is underdeveloped. Later, Bunge transferred Kochia monticola into Panderia based on the vertical embryo position. Sukhorukov et al. (2016) observed the specimens at the fruiting stage (Lebanon, 2 IX 1931, A. Eig, M. Zohary (HUJ); Lebanon, 1 VIII 1946, P. Mouterde, № 8640 (G)) and confirmed the vertical embryo position in all seeds.
Habitats. Patchy subalpine and alpine meadows, screes, roadsides at elevation 1800-2600 m a. s. l. Fl. June — September, fr. August — September.
Distribution (Fig. 1). Iran, Iraq, Syria, Lebanon.
Examined specimens. Iran. [Chaharmahal and Bakhtiari Prov.] in alpe Kuh-Delu [Kuh-e Do Delu, Kuh-e
Morphology, nomenclature and distribution of Bassia montícola (Chenopodiaceae-Amaranthaceae)
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Fig. 1. Distribution of Bassia monticola based on the specimens seen. A — distribution in Lebanon and Syria.
Dowdülü], 15 VI 1842, Th. Kotschy, № 21 (BM); Persia [Mazandaran Prov.], ad radices m. Demawend [Damavand] pr. p. Lar, 3 VIII 1843, Th. Kotschy, № 618 (BM, E, FI, G, K, LE); [Esfahan Prov.] 39 km S of Damaneh (Kumitak), bord de la route, 6 VII 1959, alt. 2140 m, H. Pabot, № 2127 (G); [Fars Prov.] Kuh-e Rang [Kuh-e Ranj], 6 VII 1959, alt. 2450 m, H. Pabot, № 2179 (G). Iraq. [Erbil Governorate] Makhmur, Ehrenberg, № 236 (LE). Syria. [Rif Dimashq Governorate] prope Zebdaine [Az-Zabadani] & prope Damascum, locis nudis nive derelictis supra Bludan, alt. 6000 ft [1828 m], 6 VI 1855, Th. Kotschy, № 49 (K000898881; BM; P04907852, LE); Anti-Lebanon [Rif Dimashq Governorate], E of Bloudane [Bludan], 9 VIII 1947, P. Mouterde, № 9063 (G). Lebanon. [Baalbek-Hermel Governorate] Inter Cedretum et jugum Baalbek, 6000 ft [1828 m], 30 VII 1855, Th. Kotschy, № 355 (K000898882; BM; P04907853); [North Governorate] Makmel [Mount], Ehrenberg (LE); Top of Djurd Aqrra, 8 VIII 1890, Post (BM); [North Governorate] mountain ranges between Ehden and Karnet es Souda [Qurnat as Sawda'], 2400-2500 m, summer snow region, 2 IX 1931, A. Eig, M. Zohary (HUJ); [North Governorate] in vicin. Arz El Rab [Arz El Rab] (i. e. cedretum supra Bcharreh [Bsharri]), solo calcareo, ca. 1950 m, 13 VI 1933, G. Samuelsson (K); [North Governorate] Cedars, 9 VII 1933, P. Mouterde, № 3433 (P05158984); [North Governorate] Jebel Materfe, 2000-2400 m, Berberidetum eretrae, 13 VII 1934, Botanical Dept. (HUJ); [Mount Lebanon Governorate, Sannine Mt.] Karakol Metiouhane, 1 VIII 1946, P. Mouterde,
№ 8640 (G); [Mount Lebanon Governorate], Mount Mnaitra, 1807 m, 34°04'52" N, 35°55'50" E, 10 VII 2005, R. Haber, M. Semaan, № 3518 (BEI; cited after Haber, Semann, 2007).
Bassia monticola is a well-recognisable species due to its small size, prostrate habit, filiform to lanceolate leaves, and small, barely noticeable tubercles on the fruiting perianth (Fig. 2: A). In contrast, typical specimens of B. pilosa (including Panderia turkestanica Iljin), which is also present in Western Asia, are much taller (20-60 cm) and characterised by a prominent branched main stem, oblong to ovate leaves, and a perianth with small white or yellowish wings at fruiting. Despite the fact that B. monticola was formerly considered as a synonym of B. pilosa, we cannot synonymize these species due to remarkable morphological differences. Their close relationship has not yet been confirmed with molecular data. However, two species sharing the same characteristics (the kochioid type of C4 kranz-type leaf anatomy and an unusual vertical seed embryo), B. pilosa and B. lasiantha Freitag et G. Kadereit (= Kirilowia eriantha Bunge), form a separate clade within Bassia (Akhani, Khoshravesh, 2013). Bassia monticola is characterised by the same seed embryo position (Sukhoru-kov et al., 2016) and leaf anatomy, and these facts support the close relationship of these three species.
The distribution area of B. monticola presented here (Fig. 1) is much smaller than that of B. pilosa and has a significant gap between central Iran and Lebanon/ Syria due to the absence of suitable habitats in the low-
Fig. 2. Perianths of Bassia monticola and Camphorosma monandra (SEM).
A — Bassia monticola (origin of the material: Lebanon, Ehden and Karnet es Souda, 2400-2500 m, 2 IX 1931, A. Eig, M. Zo-hary (HUJ)); B — Camphorosma monandra (origin of the material: Afghanistan, prov. Bamyan, Koh-i-Baba, Koh-e Fu-ladi SW Bamian, Fuladi-Kar, 3900 m, Wegrand, 5 IX 1968, № 1581, S. W. Breckle (MSB120718)).
land deserts of Syria and Iraq. Bassia monticola may also be found in Western Iran and Southern Turkey. In contrast, B. pilosa is widely distributed in the entire Irano-Turanian floristic region (Iljin, 1936, as Pande-ria pilosa and P. turkestanica; Mouterde, 1966; Aellen, 1967; Hedge, 1997, 2001, all as P. pilosa; Freitag, Ka-dereit, 2014; Sukhorukov, pers. obs. in West China and East Kazakhstan), and it prefers saline and clayey soils in lowland deserts and screes at lower elevations (up to 1500 m a. s. l.).
The southernmost record of B. pilosa is known from the Moav floristic subdivision, Jordan (Palaestina Moabitica, Mashita, deserts, alt. 750 m, 25 IV 1911, F. S. Meyers, J. E. Dinsmore, № 1658 (G!, K!)). The robust specimens turn black upon drying and are mostly known from the deserts of Syria and Southern Iraq. These specimens may belong to an undescribed species labelled in some herbaria as "Panderia iraqensis Suk-hor." (BM!, K!) or "Halostigmaria maris-mortui Eig" (HUJ!); the latter name is based on an accidental record from the northern part of the Dead Sea. The plants distributed in the deserts of Western Asia need further investigations. However, a recent record of P. pilosa from Mount Mnaitra, Lebanon (Haber, Semaan, 2007), clearly belongs to B. monticola and is in fact not the first record of this species for the country.
Another poorly known taxon, Camphorosma mo-nandra, was described by Bunge in Boissier (1879, as "monandrum") and closely resembles B. monticola in its small, prostrate and hirsute habit, linear leaves, leafy inflorescences, concrescent perianth segments, and vertical seed embryo. It is confined to the alpine belt of Central Afghanistan (in the provinces Bamyan and Maidan Wardak) and is present in several herbaria (G!, LE!, MSB!). However, it was not mentioned in recent checklists of Afghan flora (Breckle, Rafiqpoor, 2010; Breckle et al., 2013). It seems to be absent in adjacent countries (Hedge, 1997; Sukhorukov et al., 2019), and the distribution areas of B. monticola and C. monandra do not overlap. Camphorosma monandra is distinguished from B. monticola by a cup-shaped tube (Fig. 2: B) (vs. cylindrical perianth tube: Fig. 2: A) and four perianth teeth that are unchanged in fruiting (vs. five teeth with tubercles). The improved diagnostic keys to Bassia species in Western Asia were provided by Akhani and Khoshravesh (2013) and Sukhorukov et al. (2016), and B. monticola should be added to the list of Bassia species in Western Asia.
Acknowledgements
We thank Alexander Sennikov, Irina V. Sokolova, Helmut Freitag and an anonymous reviewer for useful comments to the first draft of the paper. The
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investigation was carried out with financial support from the Russian Foundation for Basic Research (grant 18-04-00029-a). The revision of the herbaria in Moscow was supported by the scientific programme AAAA-A16-116021660045-2 of the Department of Higher Plants, Lomonosov Moscow State University and by a Moscow State University (MSU) Grant for Leading Scientific Schools "Depository of the Living Systems" in the framework of the MSU Development Programme.
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