Taxonomic revision of Chenopodiaceae in Himalaya and Tibet

Abstract The composition of many Chenopodiaceae genera in different parts of Himalaya and Tibet has been insufficiently known or contradictory. A revision of the family in Himalaya including Bhutan, Nepal, parts of India (Himachal Pradesh, Jammu and Kashmir, Sikkim and Uttarakhand) and Tibet (Xizang, China) is presented for the first time. Altogether, 57 species from 20 genera are reported, including three species new to science (Agriophyllumtibeticum, Salsolaaustrotibetica and Salsolahartmannii). Atriplexcentralasiatica, Corispermumdutreuilii and Salsolamonoptera are identified as new records for India and Chenopodiumpamiricum is recorded in China for the first time. Dysphaniaambrosioides and Sympegmaregelii are recorded for Xizang. The generic and species keys, species distributions (including maps) and taxonomic notes are provided. We indicate for the first time that the presence of short yellow hairs is the remarkable morphological characteristic of the genus Grubovia. Evident heterocarpy and heterospermy is found in Dysphania for the first time (Dysphaniatibetica). Agriophyllumpungens, Atriplexcrassifolia, Atriplexlaciniata, Atriplexsagittata, Axyrisamaranthoides, Axyrishybrida, Bassiaindica, Corispermumkorovinii, Dysphaniaschraderiana (=Chenopodiumfoetidum auct.), Halocharisviolacea and Suaedamicrosperma are excluded from the species list. Neobotrydiumcorniculatum is synonymised with Dysphaniakitiae, Neobotrydiumlongii with Dysphaniahimalaica and Neobotrydiumornithopodum seems to be conspecific with Dysphanianepalensis. Corispermumladakhianum is a new synonym of Corispermumtibeticum. Amaranthusdiandrus is added to the synonyms of Acroglochinpersicarioides, and Bassiafiedleri, previously considered as conspecific with Gruboviadasyphylla, is added to the synonymy of Bassiascoparia. Lectotypes of Anabasisglomerata (≡Halogetonglomeratus), Halogetontibeticus (=Halogetonglomeratus), Amaranthusdiandrus (=Acroglochinpersicarioides), Chenopodiumtibeticum (≡Dysphaniatibetica), Corispermumdutreuilii, Corispermumfalcatum, Corispermumlhasaense, Corispermumpamiricumvar.pilocarpum (=Corispermumgelidum, syn. nov.), Corispermumtibeticum, Kochiaindica(≡Bassiaindica), Kochiaodontoptera (≡Bassiaodontoptera) and Salsolamonoptera are selected. Out of 53 native elements, 42 are restricted in their distribution to Himalaya and Tibet at altitudes 2000–4500 m above sea level. The greatest taxonomic diversity of the Chenopodiaceae is represented in Jammu and Kashmir (India) and Xizang (China) with a continuous decrease in the number of species southwards.


Introduction
The family Chenopodiaceae is one of the most difficult groups of flowering plants in terms of its taxonomy and diagnostics. In the last two decades, our knowledge of the relationships in the family, its placement within the order Caryophyllales and the physiological or anatomical peculiarities of its species have greatly increased. However, there have only been a limited number of taxonomic revisions of the family based on recent taxonomy and critical analysis of collections.

Taxonomy and generic delimitation
The Chenopodiaceae is a part of the large Chenopodiaceae-Amaranthaceae clade within the core order of Caryophyllales and it is a sister group to the Amaranthaceae family (Cuénoud et al. 2002, Kadereit et al. 2003, Brockington et al. 2009). Although all morphological characters seem to overlap in Chenopodiaceae and Amaranthaceae s. str., the familial status of the Chenopodiaceae is accepted in recent taxonomic treatments (e.g. Sukhorukov 2014, Hernández-Ledesma et al. 2015. The carpological differences that have been discovered (Sukhorukov et al. 2015a) have filled the gaps in the character set of both Chenopodiaceae and Amaranthaceae s. str., but they are not strongly specific at the familial level except the horizontal embryo position which evolved in many Chenopodiaceae.
When separated from Amaranthaceae, the Chenopodiaceae includes ~100 genera and 1600 species that are mostly distributed in the deserts and steppes in the temperate regions of the world (Sukhorukov 2014). During the last fifteen years, the generic composition of many groups in the family has undergone extensive changes following molecular phylogenetic revisions, with the splitting or merging of many genera. Chenopodium was found to be polyphyletic and its members occupy different phylogenetic positions within Chenopodioideae (Fuentes et al. 2012;Sukhorukov et al. 2018). In the previous broader sense, Salsola appeared to be strongly polyphyletic (Akhani et al. 2007), but in contrast, the formerly accepted genus Kochia is now merged with Bassia (Kadereit & Freitag, 2011). The monophyly of many other traditional genera, e.g. Agriophyllum (Kadereit et al., 2003), Suaeda (Schütze et al., 2003), Atriplex except Halimione (Kadereit et al., 2010), Corispermum (Xue & Zhang, 2011) and Dysphania, including Neobotrydium and Ambrina (Uotila et al. in prep.) is confirmed.
Subsequently, the generic status of some groups in Tibet and Himalaya has been changed compared with previous taxonomy accepted in many floristic works including comprehensive treatments of the family in adjacent territories like "Flora Iranica" (Rechinger 1997), "Flora of Pakistan" (Ali and Qaiser 2001), as well as "Flora of China" (Wu and Raven 2003) that emcompasses a part of the territory under study. Of all the known Chenopodiaceae species in the study area, only the position of Haloxylon thomsonii is still pending within the tribe Salsoleae and its placement with tree-like species within the Saharo-Arabian and Irano-Turanian genus Haloxylon may be incorrect due to differences in life history and stem anatomy (Sukhorukov, unpubl.).

Study premise
The most recent taxonomic revision of the Chenopodiaceae was performed for Nepal , but the species composition of some genera and their distributions are still insufficiently known in other parts of the entire Himalayas and Tibet. The latest Chenopodiaceae treatment for India (Paul 2012) contained many taxa that are, in fact, distributed only in Central Asia (e.g. Atriplex crassifolia, Axyris amaranthoides, Chenopodiastrum hybridum and Suaeda microsperma), Eastern Europe/North Kazakhstan (Atriplex sagittata) or the Mediterranean (Atriplex rosea). Taxonomic novelties to the chenopodiaceous flora of China were provided by Zhu and Sanderson (2017) and some new species (i.e. Neobotrydium corniculatum, N. longii) were also reported from Xizang. All these taxa reported from India or China are discussed in our article.
The present revision is a part of the treatment of the family in the Old World. Until now, the Chenopodiaceae had been revised for the flora of European Russia (Sukhorukov 2014), Iraq (Sukhorukov et al. 2016) and historical Palestine (Danin et al. in press). In addition to Himalaya and Tibet, further contributions are planned for all of Africa, Russia and Uzbekistan.

Choosing the study territory
Geographically, the Himalayan Mountains and the Tibetan Plateau (sometimes called Pan-Himalaya) encompass a large territory bordered by the Karakoram Range to the north, the Indian plate to the west, the Central China plain to the east and Mainland Southeast Asia to the south. The territory included in the present investigation ( Fig. 1) covers most parts of Himalaya and the Tibetan Plateau within the following countries: Bhutan, Nepal, parts of China (Xizang or the Tibet Autonomous Region) and India (states of Uttarakhand, Himachal Pradesh, Sikkim and Jammu and Kashmir, excluding the politically disputed territory in North Jammu and Kashmir with a poorly investigated flora that is probably rich in Central Asian species). Arunachal Pradesh (India) is not included in this treatment due to an absence of specimens seen. This treatment is a major contribution to Chenopodiaceae under the project "Flora of Pan-Himalaya".

Materials investigated
The field investigations were conducted by Alexander Sukhorukov in different provinces of Nepal in 2005 and 2008-2015 and in Dehradun and its surroundings in 2018 (Uttarakhand, India). The main collection is in MW with duplicates in B, BM, BR, E, G, H, K, M and W. The revision of the herbarium material was undertaken by the first author (AS) at B, BM, BR, BSD, DD, E, G, H, HUJ, K, KATH, L (including U and WAG), LE, LY, M, MHA, MSB, MW, P, PE, PRA, SHI, TO, TUCH, W, WU, WUK, XIA and XJBI (herbarium abbreviations according to Thiers 2018+) and the Axyris and many Dysphania specimens were revised by the first author in CAL in 2011. The Chinese Virtual Herbarium (CVH, http://www.cvh.ac.cn/) and the herbarium database of the Kunming Institute of Botany of the Chinese Academy of Sciences (Kingdonia, http://db.kun.ac.cn/) were used as citations for some specimens kept in HNWP, NAS and KUN if their identification were possible using the images. The great number of examined specimens was mostly provided by the following collectors or collection teams.
Many interesting facts about the lives of these brothers, their relationships with other botanists and the destiny of their herbarium can be found in the book by von Brescius et al. (2015). The examined collections are available from the year 1856 in the BM, E, G and LE herbaria. Thomas T. Thomson. Expeditions to West Himalaya, Tibet and then to Sikkim (Woodward 1898). Chenopodiaceae collections are available from 1845 and cluded Wang Jin-Ting, Lang Kai-Yong, Ma Cheng-Gong and Bao Xian-Cheng (Wang et al. 2004). Qinghai-Tibet Complex Expedition Team, abbreviated Qinghai-Tibet Team. Collected throughout Xizang in 1973. The team also consisted of several subdivisions, such as the "Vegetation Group", "Meadow Group" and "Additional Group". In total, nearly 40000 collections on approximately 150000 sheets were made and these specimens were deposited in PE, KUN and HNWP. Collectors worked in the Team were Wu Su-Gong, Ni Zhi-Cheng, Lang Kai-Yong, Chen Shu-Kun, He Guan-Fu, Cheng Shu-Zhi, Gu Li-Min, Nan Yong, Luosangxi'nao, Xiao Yonghui, Yang Yong-Chang, Huang Rong-Fu, Tao De-Ding, Zang Mu, Yin Wen-Qing, Su Zhi-Yun, Wu Zheng-Yi, Du Qing, Yang Chong-Ren, Guan Kai-Yun, Chen Wei-Lie, Li Bo-Sheng and Wang Jin-Ting (Wang et al. 2004).

Preparation of the material for carpological studies
The perianth, fruit and seed characters play significant roles in the identification of some large genera, e.g. Atriplex, Chenopodium, Corispermum and Dysphania (e.g. Uotila 2001, Sukhorukov 2006, 2007a, 2007b and we used scanning electron microscopy (SEM) to show the important differences in the reproductive traits. The hard tissues (seed coat) do not require any special preparation prior to SEM, but other objects that contain living cells, such as hairs or papillae (on perianth or pericarp), were dehydrated in aqueous ethyl alcohol solutions of increasing concentration, then in alcohol-acetone solutions and pure acetone and then finally critical-point dried. After sputter coating the material with gold-palladium, all SEM observations were made with a JSM-6380 microscope (JEOL Ltd., Japan) in the Laboratory of Electron Microscopy of Moscow State University.
The anatomy of the fruit and seed is important for the diagnostics and taxonomy of many Chenopodiaceae (for more see, e.g. Sukhorukov 2008, 2015a. The fruit anatomy appears crucial to the diagnostics of many Corispermum and the fruits of almost all members were anatomically investigated in previous studies (Sukhorukov 2007a, 2007b). Here, we present the fruit anatomy of Corispermum tibeticum Iljin and add the most important data to the description of the species.
The list of specimens used for SEM is given below. The altitudes from 2200 to 3500 m above sea level. Most annual Chenopodiaceae growing at these altitudes are found on the screes, on limestone or sand or often in disturbed habitats (being apophytic). In North Himalaya and Tibet, some annuals that usually grow at 3500-5000 m above sea level can be found at lower altitudes (Chenopodium sp. div., Corispermum sp. div. and Dysphania spp.). In Central Himalaya, 2200-3500 m is the most appropriate altitude for Acroglochin persicarioides and Dysphania nepalensis, where they are abundant, weedy species . In West Nepal, Chenopodium perttii and Dysphania neglecta are common. Different Chenopodium species often grow together, e.g. Dysphania neglecta and D. nepalensis in Nepal (Fig. 3).
The altitudes from 3500 to 5000(5500) m above sea level. This altitude range constitutes a zone of alpine steppes and cold deserts (Fig. 2) where three sub-shrubby Chenopodiaceae members play a significant role as the dominants of different plant communities. Krascheninnikovia ceratoides (s.l.) is one of the dominants of stony plant formations of North Himalaya (Hartmann 1983(Hartmann , 2009) and Tibet (Namgail et al. 2012, Hong andBlackmore 2015), with local distribution in Nepal, Central Himalaya (Yonekura 2008, Sukhorukov and. Other subshrubs have limited distribution patterns in the territory under study. Bassia prostrata is mostly distributed in Jammu and Kashmir (India), where it can be abundant in subalpine deserts at altitudes of 3500-4000 m (Hartmann 2009). Another dwarf subshrub, Haloxylon thomsonii, is localised only in Ladakh and adjacent territories in Pakistan at altitudes of (3200)3500-4000 m, where it forms characteristic alpine semi-desert and steppe landscapes together with some Artemisia, Caragana, Potentilla and Ptilotrichium species (Dhar and Kachroo 1983, Hartmann 1983, Gruber and Peer 2008. Sympegma regelii, a common plant on the screes in Tian-Shan and Kunlun ), has been found in northwest Tibet during the last several decades.
Amongst the annuals, Axyris mira, Axyris prostrata, Corispermum sp. div. and some Dysphania (D. himalaica, D. nepalensis and D. tibetica) are widespread in stony deserts, river basins and disturbed areas. Microgynoecium tibeticum, a small, often-overlooked herb, reaches altitudes up to 5500 m, and this altitudinal zone appears to be the highest level at which only a few flowering plants are observed (Sukhorukov 2014). All Chenopodiaceae at the higher altitudes are native species growing in natural or degraded communities (Klimeš andDickoré 2005, Miehe et al. 2009).

Taxonomy and geographical subdivision of the species
In their recent circumscriptions, 20 genera are recognised in Himalaya and West Tibet, representing all subfamilies of Chenopodiaceae and 57 species have been recorded. The best-represented genera are Chenopodium, Corispermum and Dysphania (9 species each). None of the taxa can be recommended for inclusion in the IUCN Red List (IUCN 2017) since they are either dominant species, occupy ruderal habitats or are insufficiently studied.
In the past, very few species were recorded for the entirety of Himalaya and Tibet (within the borders of Xizang) and the number of Chenopodiaceae species reported for Tibet was only 15  or 29 (Zhu et al. 2003). Fifteen species were reported for Lower Ladakh, India (Klimeš and Dickoré 2005). Here, we present the first attempt to evaluate the number of species in the main divisions of the region under study. The number of species was counted for all the countries of Bhutan and Nepal, the major administrative subdivisions of Xizang (prefectures) and the Indian states of Jammu and Kashmir, Himachal Pradesh, Uttarakhand and Sikkim (Fig. 1).
Two species, Atriplex hortensis and Dysphania ambrosioides, are adventive plants (ergasyophytes). The adventive status of Chenopodium album and C. ficifolium is still not confirmed.

Key to the genera
The key to the infrafamilial groups (subfamilies) and their characteristics are provided in some recent works (Sukhorukov 2014;Sukhorukov et al. 2015aSukhorukov et al. , 2016b) and we chose not to repeat it in the present work. The generic key does not include Spinacia oleracea L. (Chenopodioideae-Anserineae), which is often cultivated as a vegetable (especially in China) and Beta vulgaris L. (Betoideae), which is also sometimes present in vegetable gardens. Both species have no tendency to escape from cultivation. A formal proposal to conserve the name Chenopodium L. with the type Chenopodium album L. has only recently been put forward (Mosyakin 2015). The conservation of the name with this type would be most desirable to maintain the recent taxonomy of Chenopodium and its relatives, especially Oxybasis.
Description. Annuals, shrubs, or rarely small trees, covered with bladder hairs. Leaves petiolate, usually lobed or dentate (sometimes entire), very rarely semi-terete. Inflorescences paniculate, composed of small cymose clusters. Flowers sessile and pedicellate, hermaphrodite or some female. Perianth segments 5 (rarely 4), free or basally concrescent, green and unchanging at fruiting. Stamens usually 5, free or basally connate. Stigmas 2, free. Fruit depressed globular, falling off separately or together with perianth. Pericarp mostly thin, hyaline, of 1-2(3) parenchymatous layers, usually with small cylindrical or conical papillae (in dry fruits, the pericarp surface resembles honeycombs, but after soaking, the papillae retrieve their shape). In some species now transferred from Einadia to Chenopodium, the pericarp (at least in most fruits) is fleshy (berry), coloured, and many-layered, but some of the fruits remain dry (heterocarpy). Seeds black or rarely brown with horizontal embryo; the cell walls of testa (outer seedcoat layer) cells in black seeds with vertical stalactites, rarely the stalactites absent (three species from Juan-Fernández Archipelago).
In its current circumscription (Fuentes et al. 2012), the genus comprises more than 100 species, but the exact number is still unknown. The species are mostly distributed in extratropical parts of the world or in mountainous regions of the tropics.
Key to the species vulvaria, but all data on the presence of C. vulvaria in mountainous regions of Central Asia belong to C. pamiricum. In contrast to the latter species, either fresh or dry leaves of C. pamiricum do not have a scent. Further investigations have shown that the type material and many other specimens of C. pamiricum have evident heterospermy, a rare characteristic in Chenopodium s. str. discovered by .
Description. Annual up to 25 cm with several or numerous prostrate or ascending stems. Leaves farinose, shortly petiolate (petioles up to 1.5 cm but usually much smaller) with blades up to 2 cm, rhombic or ovate, entire or subhastate. Inflorescence leafy almost to the top. Flowers in glomerules arranged in ± dense inflorescence. Perianth farinose. Fruits and seeds heteromorphic. The first type of fruits and seeds is formed in terminal flowers, the fruits ripen in July-August. These fruits are ca. 1 mm in diameter and 0.5 mm thick; pericarp with papillae up to 20(25) µm (Fig. 4C); seed dark red or blackish with median keel (Fig. 4E, F), testa 17-25 µm thick. Fruits of the second type (which dominates at the end of the vegetation period) are ovoid, 1.0-1.2 mm long, ca. 0.5 mm thick, pericarp papillate or not (Fig. 4D); seeds yellowbrownish, elongated.
Habitat. Sandy or rocky places, 3400-5000 m a.s.l. (in Himalaya) and at lower altitudes in Central Asia. General distribution. Central Asia south to North Himalaya and North Tibet. For a long time, C. pamiricum has been considered endemic to Pamir (Iljin 1936, Pratov 1972. The analysis of the material collected in the fruiting stage proved that the range of Chenopodium pamiricum covers almost all parts of Central Asia with records in Mongolia, West China (Xizang and Xinjiang), North Pakistan, North India, Kyrghyzstan, Tajikistan, Russia (Altay Mountains) and Eastern Kazakhstan. The records from Pamir, Karakoram, NW Tibet and North Himalaya represent the southernmost boundary of the distribution of C. pamiricum.  (1800) is probably lost. The specimen col- lected in England [Yarmouth, Sep 1801, Mr. D. Turner] and kept at LINN-HS (sheet 459.24) is not a part of the original material used for the description. Citing the "type", Al-Turki and Ghafoor (1996) refer to the drawing of Curtis (1778, fig. 16), but Curtis did not describe any Chenopodium in his work and erroneously labelled his fig. 16 as "Chenopodium viride", although the image of the plant completely corresponds with Chenopodium ficifolium later described by Smith (1800). Citing the type of Chenopodium ficifolium, Al-Turki and Ghafoor (1996) have unintentionally lectotypified the species with the drawing in Curtis (1778). The chosen lectotype shows the plant with clearly trilobate leaves with prominent and elongated middle lobe (Chenopodium ficifolium subsp. ficifolium: Uotila 1997Uotila , 2001.
Habitat. Disturbed places, river-sides, sands; 0-2200(3000) m a.s.l. Common at least in many parts of Nepal and Bhutan. Numerous seedlings were observed by AS near Mussoorie (Uttarakhand, India) in mid-February 2018.
Phenology. Flowering: February-May; fruiting: April-June. At higher altitudes (where the plant is rare), the blooming and fruiting times appear to be much later.
Distribution. See Fig. 6 rukov and Kushunina 2014), this species is cited as Chenopodium pallidum Moq., but a closer look at the authentic specimens in P has proved that all plants (collected in early blooming stage) are indeed Atriplex pallida, now synonymised with A. schugnanica Iljin .
Description. Annual up to 40 cm with erect stem; lateral branches ascending if present. Leaves short-petiolate (petioles to 1 cm) with blades up to 2 × 1 cm, fleshy, ovate, slightly cuneate, green above, greenish below, dentate, margins often red. Inflorescence bracteate or aphyllous. Perianth green, enclosing the fruit; segments abaxially carinate. Fruit 1.3-1.5 mm, pericarp easily separating from the seed; papillae up to 25 µm. Seed blackish or reddish-black, surface striate with obtuse or acutish margin (immature seeds with acute margin) with small depression apically.     (Uotila and Lomonosova 2016). However, C. jenissejense is quite distinct in its morphological characters and needs to be studied in detail.

Description.
The species is similar to C. album, but plants are dark green, usually with ascendant or prostrate stems up to 25 cm. Leaves ± ovate, somewhat longer than broad, sometimes inconspicuously three-lobed.  Lo Tsho Dhyun, Yara [Yara Gaun] (1840). Description. Robust annual up to 2.5 m, branched from the base. Petioles of lower caducous leaves up to 10 cm, blades 10-25(30) × 5-10 cm, usually rhombic, dentate, erose or trilobate with elongated terminal lobe; middle leaves rhombic or ovate, smaller; upper leaves oblong or lanceolate, completely entire, abaxially grey; young leaves often reddish. Inflorescence whitish due to the presence of abundant bladder hairs on the branches and perianth. Fruit 1.2-1.5 mm, pericarp hyaline, adjoining to the seed coat, but separating from it when rubbed, long-papillate in fresh or even dry fruits (Fig. 8D). Seed black, ± keeled, with small crater-like micro-depressions (Fig. 8E, F).
Habitat. Edges of vegetable gardens and other places with secondary vegetation; 700-2900 m a.s.l. Cultivated as a crop but very locally.
Taxonomic notes. This species was described from the deserts of Turkmenistan and the identity of the populations growing in the Central Asian plains and in the Himalayan Mountains is not obvious despite the morphological similarities. The material from Himalaya is scarce and collected in the blooming or early fruiting stages. Further investigations are needed to establish the taxonomic status of Himalayan plants named C. novopokrovskyanum. Description. The species is similar to C. album, but the leaves are bi-coloured (green or greyish adaxially and white-farinose abaxially), narrowly oblong to lanceolate. Fruit 1.0-1.1 mm in diameter, ca. 0.6 mm thick, seed almost smooth with indistinctly polygonal ultra-sculpture and indistinct pits (Fig. 10A Lectotype (Uotila 1993): AFGHANISTAN, Kurrum valley, 1879, J.E.T. Aitchinson 980 (G11875! isolectotypes BM000629112! C, G-BOISS! FI, K! S). Taxonomic notes. Murr (1902) cited two distant records (Russian Far East and Afghanistan) in the protologue of C. atripliciforme. We agree with Uotila (1993) that the specimen from the first location (Ussuri) probably belongs to C. bryoniifolium Bunge, which is similar in leaf shape to C. atripliciforme.
Description. Annual up to 1 m, mostly branched; leaves petiolate (petioles up to 4 cm). Leaf blades triangular, basally truncate or broadly cuneate, entire to erosedentate. Inflorescence branched, terminal and axillar, slender; clusters consisting of 2-4 flowers or rarely flowers solitary. Fruit 1.2-1.4 mm with papillate pericarp. Seed black, almost smooth with indistinct small pits (Fig. 10C, D).  Taxonomic notes. Morphologically, Chenopodium perttii resembles a Far Eastern and Siberian species, C. bryoniifolium Bunge, but the species have different distributions and reproductive characters . The recently identified chromosome number for C. perttii is 54, so it is genetically distant from other species previously identified as C. album (Bohumil Mandak, pers. comm.). However, we assume that C. perttii is more closely related to Chenopodium album subsp. yunnanense Aellen in Handel-Mazzetti (1929). This subspecies was described from Yunnan (China) and differs by the absence of the seed keel. The lectotype of Chenopodium album subsp. yunnanense was selected in .
Description. Annual up to 1(1.5-2) m with erect, scarcely branched stem. Leaves dark green, petiolate (petioles up to 3 cm), blades 3-4 × 2-3 cm, base truncate or broadly cuneate, margins often red; lower leaves triangular, entire to lobate; middle leaves trilobate with elongated mid-lobe (less than two times as long as the lateral ones) gradually tapering to apex and slightly upward-directed lateral lobes, lobes mostly entire; upper leaves also trilobate or entire, narrowly oblong. Inflorescence green, aphyllous or rather bracteose. Fruit ~1.5 mm, pericarp separating from the seed (not easily) with minute papillae that are hardly restored after soaking, almost smooth in dry fruit. Seed prominently keeled, surface with radial striae ( This name was previously widely used for the plants growing in Europe, Siberia, Iran and Central Asia (e.g. Iljin 1936, Kinzikaeva 1968, Lomonosova 1992, Uotila 1997. Sukhorukov (2014) discovered that the plants growing in North Himalaya and in Europe differ in their morphological characters and C. betaceum Andrz. may be the correct name for the European plants. Furthermore, C. strictum material from Himalaya, from where the species was described, is scarce (Jammu and Kashmir, India). Further investigations are needed to discover which plants belong under this name.

Chenopodium hookerianum Moq. in DC., Prodr. 13(2): 68 (1849)
Notes. The species was described from Nepal, ex herb. Hooker (P00606414) (see link to the specimen https://science.mnhn.fr/institution/mnhn/collection/p/item/ p00606414) and the type material bears a plant fragment collected at the flowering stage. This name has never been mentioned in the recent floristic and taxonomic accounts except the Flora of the USSR (Iljin 1936), where it was synonymised with Chenopodium giganteum (=C. bengalense). However, the protologue of C. hookerianum (Moquin-Tandon 1849) and a fragment of the type specimen do not demonstrate the features of the species described by Moquin well (plant height, seed characters etc.). We are still not sure how to treat C. hookerianum.

Chenopodium patulum Roth, Nov. Pl. Sp.: 180 (1821)
Notes. The species was described from "India Orientalis" [East India] without a precise location. The type material was not found in any herbaria including HAL. The name is a later homonym of C. patulum Mérat described from France (Mérat 1812) and cannot be used according to ICN (Art. 53.1).
Description. Annuals, glabrous or covered with bladder hairs. Leaves triangular or rhombic, petiolate, entire, dentate or lobate, rarely pinnatisect. Inflorescences spreading, mostly leafless, often with scattered yellowish glandular hairs. Flowers bisexual or female with 5 perianth segments that do not change at the fruiting stage. Perianth segments free or connate to the half of their length; midvein clearly elevated adaxially. Stamens 5. Stigmas 2. Fruit 1.2-2.5 mm in diameter; pericarp 1-2-layered with conical or cylindrical papillae, forming alveolate (in fresh fruits) or reticulate (in dry fruits) surface. Seed black, with keel or not, alveolate or punctate, sometimes with deep combs; seed-coat testa with vertical or obliquely orientated stalactites. Latent structural heterospermy expressed as varying testa thickness found in some species; seed embryo horizontal.
At least 10 species in all continents.
Key to the species Taxonomic notes. Chenopodium ilicifolium was previously synonymised with Chenopodium botrys (≡Dysphania botrys); see Press et al. (2000). Despite the absence of the authentic herbarium, we merge Chenopodium ilicifolium with Chenopodiastrum murale for the following reasons: (1) the drawings of C. ilicifolium (Griffith 1854) show the plant with dentate leaves, a short and leafy inflorescence, closed flowers and fruit with a papillate pericarp. These traits are characteristic of C. murale. Additionally, the city of Shikarpur is located on the plains where other Chenopodiastrum do not occur. The merger of Chenopodium ilicifolium with Chenopodiastrum murale is consistent with the opinion of Singh (2014, sub Chenopodium murale). Description. Annual, 10-50 cm, erect, with several lateral branches, slightly mealy. Leaves 2.0-6.0 × 1.0-5.0 cm, dark green, broadly deltoid-ovate, upper leaves narrow and long-acuminate, mucronulate, cuneate, coarsely and irregularly dentate, rarely subentire or erose-dentate; teeth acute and ± incurved. Inflorescences terminal and axillary (leafy) with loosely branched cymes. Perianth segments slightly keeled with a distinct swelling below the apex. Fruit 1.3-1.4 mm, pericarp tightly adjoining the seed, papillate. Seed with prominent keel and surface densely covered with small pits (Fig. 13A   General distribution. Widely distributed in tropics and subtropics in all parts of the world. Origin unknown, but Sukhorukov (2014) proposed that it may be Eastern Africa and Arabia, where the morphologically similar C. fasciculosum (Aellen) Mosyakin occurs. Chenopodiastrum murale is rather common in the plains of India, but there are limited collections from the Himalayan foothills. The species was not observed by AS in Dehradun (Uttarakhand, India) and surroundings in February 2018. Taxonomic notes. The "true" Chenopodiastrum hybridum reported for some territories (e.g. Chowdhery 1984, Zhu et al. 2003 is not present in Himalaya and Tibet. Additionally, we are uncertain that C. badachschanicum described from the Pamir Mountains (Tzvelev 1960) is the correct name for our populations. As recently recognised (Sukhorukov in prep.), the seed surface plays a significant role in the diagnostics of many species of the genus. The type material of C. badachschanicum bears small plants (up to 25 cm) with acutely lobed leaves but without ripe seeds. The similar individuals at the fruiting stage were collected in East Kazakhstan, Kyrgyzstan and Tajikistan and their fruits have adherent pericarp and clearly pitted seeds (like those of C. hybridum). The plants growing in Himalaya and Tibet are quite uniform in their morphological characters and possess the fruit with loose pericarp and rugulose seed surface. C. badachschanicum complex needs further investigations.
Description. Annuals, branched from the base or with a single stem, prostrate or erect, glabrous or covered with bladder hairs. Leaves alternate, entire to lobate, rhombic, triangular or oblong, sometimes lanceolate, green from both sides or whitish abaxially. Inflorescence racemiform with lateral branches mostly appressed to the stem; flowers arranged in dense glomerules. Perianth of 3-5 free or diversely connate, hyaline or greenish segments (both perianth forms are present in some species), keeled or not. Flowers bisexual or sometimes female (lateral flowers). Stamens 2-5. Stylodia 2. Pericarp thin, smooth, mamillate or rarely papillate. Seeds usually small (up to 1.2 mm in diameter), red or black. Embryo horizontal or vertical and both embryo positions may be present in one individual (spatial heterospermy). Structural (latent) heterospermy expressed in the varying thickness of the seed coat is common in almost all representatives; outer cell wall of the testa cells with stalactites. Twelve to fourteen species in the temperate and mountainous parts of Eurasia, Africa and the Americas. One species in Himalaya and Tibet. Description. Annual up to 70 cm, branched from the base, sometimes forming cushion-like habit at high elevations; stem often prostrate to ascending, rarely straight. Leaves up to 6 × 2 cm, petiolate, cuneate at base, oblong or lanceolate, dentate or lobate with 2-5 lobes, rarely entire, green adaxially, grey or whitish below. Inflorescence leafy, loose. Perianth segments 3 to 5, almost free with hyaline margins, keeled along midrib, opened at fruiting stage. Fruit 0.65-0.8 mm in diameter, pericarp smooth, whitish, often ruptured. Seed reddish, without keel; embryo in both vertical and horizontal positions (spatial heterospermy), structural (latent) heterospermy expressed in different thickness of seed-coat testa.
Habitat. Rocks, streams or other wet (often sandy) places; 2800-4600 m a.s.l. Phenology. Flowering: July-September; fruiting: August-October. General distribution. Eurasia (mostly temperate regions), North America, North Africa. In Himalaya and Tibet, the species is found at high altitudes (2800-4600 m a.s.l.) and the plants hardly reach 30 cm in length. Its frequency is not known, but it seems to be rare in our area. Description. Annual herbs, subshrubs or shrubs, monoecious or dioecious, covered with bladder hairs having short basal cell and globular terminal cell; occasionally, other hair types (simple bristle-like or curved hairs) can be detected under higher magnification. Leaves alternate or opposite, simple, flat, petiolate. Inflorescence leafy or not, consisting of few or numerous flowers that are usually unisexual. The male and bisexual flowers are enclosed by (3)(4)5 green perianth segments that remain unchanged at fruiting in bisexual flowers; female flowers enclosed by two accrescent perianth segments [valves] (often called 'bracteoles' or bract-like cover) that are free or connate to varying degrees. Seeds have a vertical embryo, some seeds rarely with horizontally orientated embryo. Often (in annual species) heterospermy is present (seed coat black and brown/red).
Key to the species 1 Leaves opposite to alternate, ovoid or triangular, usually reddish and sparsely covered with bladder hairs (except young leaves, which are densely covered with bladder hairs). Plant bears three types of flowers (male and bisexual flowers surrounded by perianth of (4)5 normal segments and female flowers supported only by a bract-like cover formed of two accrescent valves that are only basally connate); seeds heteromorphic (black and brown  Pratov. We believe that these two species are indeed varieties of A. centralasiatica. In the preliminary, recently constructed ITS tree (Sukhorukov et al. in prep.), the specimens of A. centralasiatica and A. megalotheca (A. tianschanica was not included in the analysis) are intermixed within one subclade (as a part of the C 4 -Atriplex clade). A closely related Atriplex sibirica possesses bract-like cover with warty or spiny outgrowths, whereas A. centralasiatica is distinguished by having two forms of the valves, with and without outgrowths (smooth on dorsal side). The results of the first molecular phylogeny of the genus support the existence of two morphologically similar species, A. sibirica and A. centralasiatica (Kadereit et al. 2010). The distribution area of A. sibirica mostly covers the northern and middle parts of Central Asia and no specimens were seen from Tibet and Himalaya. For this reason, A. centralasiatica is still considered to be in its own specific rank.
Habitat. Screes, roadsides at altitudes of 3500-4800 m a.s.l. (in Himalaya and Tibet). Widespread in Tibet and in some parts of Ladakh (India).
Phenology. Flowering: July-September; fruiting: September-October. The records from India (Ladakh)    Taxonomic notes. Atriplex pamirica is a morphologically highly recognisable species (Sukhorukov 2006). The varietal rank Atriplex tatarica var. pamirica, proposed by Chu in Kung and Tsien (1979), cannot be accepted. Atriplex tatarica is a taller plant with an aphyllous or bracteose inflorescence. The results of molecular phylogeny have confirmed the specific rank of A. pamirica (Kadereit et al. 2010).

Atriplex pamirica
Description. Annual up to 30(50) cm, branched at the base with ascending stems but not forming tumble-weed habit at the fruiting stage. Stems and leaves covered with white and brown bladder hairs. Leaves grey on both sides, oblong or spatulate, entire or crisp. Inflorescence leafy. Bract-like cover enclosing the female flowers 3.0-7.0 mm long, rhombic, entire or dentate with concrescent (up to 1/3-1/2) valves, valves sessile or pedicellate (pedicels up to 3 mm), sclerified and inflated in the lower part, abaxial surface smooth or with two small outgrowths. Seeds heteromorphic: red seeds convex, 1.0-1.7 mm, brown seeds flattened, 1.2-1.8 mm.
Habitat. Screes or roadsides at altitudes of 3500-5300 m a.s.l. Phenology. Flowering: July-September; fruiting: September-October.  Description. Annual up to 150 cm, erect, usually branched. Leaf petioles 1.5-4.0 cm, blades 4.0-15.0(22.0) × 3.5-7.0(8.0-20.0) cm, triangular-hastate or ovoid, entire or dentate, basally slightly cordate or rounded, apically obtuse, green or reddish, glabrous or with scattered bladder hairs abaxially. Inflorescence mostly basally leafy, spike-like. As a rule, three kinds of flowers are present: (i) male flowers surrounded by a perianth with (4)5 segments, (ii) bisexual flowers surrounded by a perianth with (4)5 segments (sometimes not observed in the Indian specimens) and (iii) female flowers supported only by two accrescent, basally connate perianth valves forming a bract-like cover up to 12.0(15.0-20.0) mm in diameter, valves sessile or sometimes tapered into a peduncle up to 5 mm long (seen on Indian material). Seeds heteromorphic (three seed types present on one individual). Black seeds (1.6-1.8 mm in diameter) are formed in second (ii) and third (iii) flower types while the brownish ones (2.0-3.5 mm) are usually yielded by female (iii) flowers.
Habitat. Disturbed places (cultivated in vegetable gardens or as a weed); 2500-3600 m a.s.l. (in our area).
Phenology. Flowering: July-September; fruiting: September-October. General distribution. Eurasia (seems to be native in southern Europe and Turkey only); cultivated and sometimes well-established in Northern Himalaya, North and South America and Australia.

Excluded species
All records of the species listed below are erroneous (see for more Sukhorukov 2006) and belong to different species listed above or even to some Chenopodium.  Description. See the genus description. Microgynoecium tibeticum is often confused with Chenopodium pamiricum; both species are present in North Himalaya at high altitudes. In the early blooming stage, M. tibeticum differs by having unisexual flowers and long (2.0 mm) filaments of stamens protruding from the perianth (Chenopodium pamiricum has bisexual flowers and shorter filaments). The fruit and seed characters greatly differ between the genera.
Habitat. Screes, grassy slopes or ruderal sites; 3500-5500 m a.s.l. Phenology. Flowering: August-October; fruiting: September-November. Distribution. See Fig. 17.  Description. Annual or short-lived perennial up to 60 cm tall, glabrous or covered with scattered glandular hairs in the upper part. Basal leaves numerous, up to 25 cm long, cauline leaves up to 20 cm, long-petiolate with triangular blades, truncate or slightly cuneate at base, margins erose-dentate or lobate. Inflorescence leafy to the apex; bracts dentate to lobate. Flowers in dense clusters up to 8 mm in diameter. Perianth segments 3-5, basally concrescent, green and membranous at anthesis but usually turning red and fleshy at the fruiting stage (sometimes remaining green). Pericarp adhering to the seed coat, hyaline, consisting of 1(2) very thin layers. Seeds ovoid, 1.0-1.2 × 0.7 mm, dark red, with a groove and two blunt keels.
Phenology. Flowering: June-August; fruiting: July-September.   Description. Aromatic annuals or, rarely, perennial herbs and small subshrubs covered with glandular hairs, subsessile yellow or orange glands and simple white hairs.
Leaves alternate, entire, lobate or pinnatisect. Inflorescence lax or dense, leafy or not, consisting of cymes that are often reduced to one flower only. Perianth segments 2-5, free or variously connate, the midrib usually with keel. Stamens 1-5. Styles 2, free or basally connate. Fruit subglobose or rarely flattened, 0.3-1.5 mm, its surface reticulate or papillate, rarely smooth, dark coloured but often with whitish longitudinal stripes. Pericarp adjoining the seed coat, rarely bursting, hyaline, consisting of 1-2 very thin layers. Seed reddish or reddish-black, sometimes brown, its testa (outer seed coat layer) lacking vertical tannin-like deposits called "stalactites" in cross-section. Embryo horizontal or vertical, sometimes in both positions within one plant (spatial heterospermy). More than 50 species worldwide, mostly in the tropics and subtropics. According to the first comprehensive molecular phylogeny (Kadereit et al. in prep.), the genus Dysphania is monophyletic and there is no reason to accept the genera Neobotrydium or Ambrina as was recently proposed Zhu 2016, Zhu and. All native species are closely related and belong to a large clade consisting of Eurasian and African representatives. The mountainous areas of Himalaya and Tibet are now considered one of the four centres of Dysphania diversity, together with Australia, South America and East Tropical Africa Kushunina 2014, Sukhorukov et al. 2016a). The evident structural fruit and seed dimorphism in Dysphania (D. tibetica) is stated here for the first time.
Key to species   Description. Annual or short-lived perennial up to 1.5(2.0) m, very aromatic, covered (at least the young parts of the plant) with curved simple hairs, yellow (subsessile) glands and glandular hairs with a prominent stalk. Leaves long-petiolate, 5.0-16.0 × 1.0-4.0 cm, elliptic-oblong or lanceolate, dentate or sinuate; upper leaves often entire. Inflorescence usually highly branched, spike-like, bracteate or aphyllous in the upper part. Flowers sessile. Perianth segments (4)5, green, ca. 1.0 mm long, (nearly) half concrescent, concave near the apex, completely enclosing the fruit (Fig. 20A). Pericarp thin, hyaline, tightly adjoining the seed coat but separating from it when rubbed, in its upper part covered with glandular hairs (up to 0.12 mm long) with a large terminal cell (Fig. 20B). Seed dark red or almost black, 0.7 × 0.5-0.6 mm, not keeled. Embryo horizontal, rarely oblique or vertical.
Habitat. Disturbed places and riversides, gravelly substrates; 0-2300 m a.s.l. Phenology. Flowers and fruits year-round. Fruits develop only at elevations up to 2300 m. At higher elevations, the plants, even if occasionally present, do not produce flowers and fruits .
Habitat General distribution. South-West, Central and North-East China (for more, see ).
Habitat. Mountain steppes with Artemisia sp. div., grasslands, sands, screes or as a weed; 2000-4000 m a.s.l. Common in many parts of Jammu & Kashmir and Himachal Pradesh. Description. Annual, 5-50 cm tall, stem erect, often scarcely branched and then only in the upper part, branches directed upwards. Leaves pinnatifid or lobate, rarely toothed, appressed to the stem, up to 2.5 cm long, shortly petiolate (petioles 0.2-1.0 cm); lower leaves caducous at the flowering and fruiting stages. Inflorescence leafy, lateral branches appressed to the stem or obliquely directed, narrowly cylindrical but not spreading, axis covered with short simple hairs up to 0.3 mm long mixed with glandular trichomes. Perianth segments 5, 0.6-0.7 × 0.3-0.4 mm, cristate on dorsal surface (especially on midrib) with numerous large simple stout hairs up to 0.3 mm, the bases of which can sometimes be concrescent, resulting in 2−3-furcate tips, intermixed with yellow subsessile glands and scattered glandular hairs (Fig. 23C). Fruit subspherical, 0.5-0.6 mm in diameter; pericarp with tiny volcano-like papillae (Fig. 23D). Seed blackish, keeled, embryo horizontal.
Habitat. Grassy slopes, river banks and sands at 2500-4200(4300-4800) m a.s.l. Phenology. Flowering: July-September; fruiting: September-October. Distribution. See Fig. 24. General distribution. South Tibet, Eastern and Central Himalaya. The species can also be found in India (especially in Arunachal Pradesh) and North Myanmar.
Habitat. Gravelly slopes and ruderal sites at elevations of 3800-5000 m a.s.l. Some specimens from Ladakh contain both D. tibetica and D. himalaica, and we assume they can often grow together.
Phenology. As in D. nepalensis but fruiting 1-2 weeks later, in October (according to the observations in West Nepal).
Distribution. See Fig. 27.   (70)   Notes. This species, which is distributed in East Africa and Arabia and found as alien in many (sub)tropical areas of the world, is absent in our area. All records (Zhu et al. 2003) belong to other (native) Dysphania species, mostly D. nepalensis. Description. Annual up to 25 cm forming tumble-weed habit, non-aromatic, highly branched from the base, branches usually terminating with acicular apices, glabrous or with short papillae and white glandular hairs (mostly present in lower stem parts). Leaves up to 6.0 cm long, narrowly oblong or spatulate, sessile or with petiole-like base, entire, often folded on the ventral side. Inflorescences usually terminate with aristae, sometimes (in wet habitats) without acicular apices; flowers solitary in the axils of falsely dichotomous branches. Perianth segments 5, free at base, with slightly keeled midrib, hyaline or pinkish, glabrous. Styles 2(3). Fruit 0.7-0.8 mm, compressedly spherical. Pericarp tightly adjoining the seed coat and separating from it when rubbed, without papillae. Seeds reddish-black, keeled. Embryo horizontal, rarely obliquely or vertically orientated. One species mostly distributed in Central Asia and alien in many parts of temperate Eurasia and North America. General distribution. Central Asia, alien and ephemerophyte in many temperate regions of Eurasia and North America. In some of the herbaria visited, the specimens from Xizang identified as Chenopodium aristatum (≡Teloxys aristata) indeed belong to Dysphania species, although its wider occurrence in this province has been predicted. There are no specimens from India, but the species could be found in some areas (especially in Jammu & Kashmir).
Description. Monoecious annuals covered with stellate hairs sometimes intermixed with simple multicellular hairs. Leaves short-or long-petiolate, blade ovate, oblong, spatulate or lanceolate, entire, rarely crisp. Male flowers arranged in terminal spike-like inflorescences up to 8.0 cm long, with minute perianths of 3-5 free hyaline segments and with 2-5 stamens; female flowers located in the bract axils, with five perianth segments. Fruits always dimorphic (heterocarpous); pericarp tightly adjoining the seed coat with small ear-like appendages at the apex of the fruit. Seeds also dimorphic (with thick and thin seed-coat testa). Embryo vertical, horseshoe-shaped or annular; perisperm present.
Six species in Eurasia, predominantly in Central Asia; one (A. amaranthoides) is found as an alien plant in many parts of Europe and North America. Fruit morphology and anatomy as well as the peculiarities in plant pubescence are considered the most valuable characters for species delimitation (Sukhorukov 2005(Sukhorukov , 2011.   Sukhorukov (2005). Description. Annual with several prostrate stems up to 25 cm long; at high elevations (4000-5000 m), the plant often has a pincushion habit with small branches and short internodes. Leaves long-petiolate, spatulate, up to 3.5 cm, entire, with longer (up to 1.5 mm) stellate hairs near the leaf base, substituted in other parts by short-rayed hairs. Male inflorescence up to 2.0 cm. Perianth of the female flowers with scattered simple hairs. All fruits compressed; black fruits 1.5-2.2 mm long, oblong, with hardly noticeable or indistinct ear-like appendages; brown fruits 1.3-1.8(2.2) mm long, pearlike, with small appendages.
Phenology. Flowering: August-September; fruiting: September-October. Description. Annual up to 40 cm, main stem erect with ascending branches, green or red, densely covered with short-rayed (up to 1.0 mm) stellate hairs at least in its upper part. Leaves short-petiolate, oblong or ovate, up to 6.0 × 2.0 cm, with shortrayed stellate hairs, dark green. Male inflorescence up to 3.0 cm long. Perianth of the female flowers with dense and large simple hairs. All fruits compressed, oblong; black fruits 1.5-1.8 mm long with very small (up to 0.15 mm) appendices on the pericarp that are indistinct in most cases; brown or reddish-brown fruits (1.6)2.0-2.4 mm long.
Phenology. Flowering: August-September; fruiting: September-early November. Description. Annual up to 40 cm, erect; stem with stellate hairs having a large central ray of 0.7-2.5(3.0) mm long, such hairs especially abundant on stems and leaf petioles. Leaves short-petiolate or subsessile, up to 5 cm, oblong. Male inflorescence up to 2.0 cm long. Perianth of the female flowers with sparse, but large simple hairs. Fruits of two types: subglobose, black, 1.4-1.6 mm with hardly noticeable or indistinct earlike appendices; the second type ± compressed, ovoid, blackish, 1.6-2.0 mm, also with hardly noticeable or indistinct ear-like appendices.
General distribution. Central Asia, Siberia and Himalaya. The southernmost border of the range is in N Himalaya.

Species excluded
Axyris amaranthoides L., Sp. Pl.: 979 (1753) Notes. This species was long considered in broader sense by the authors of Himalayan floras (Hooker 1890, Hemsley 1902, Strachey and Duthie 1906, Bamber 1916) and included Axyris hybrida and A. prostrata. Subsequently all records were re-identified as A. mira or A. prostrata (Sukhorukov 2011). The southernmost records of A. amaranthoides were discovered outside of our region, e.g. in the provinces of Shaanxi, Gansu, Xinjiang in China (Sukhorukov 2014).

Axyris hybrida L., Sp. Pl.: 980 (1753)
Notes. The records of A. hybrida in Xizang and Yunnan (Zhu et al. 2003) belong to A. mira (Sukhorukov 2011), and the distribution of the former species lies in the northern part of Central Asia (including Asian Russia, Mongolia, E Kazakhstan, Kyrgyzstan, Tajikistan and N and NE China). Description. Subshrubs or shrubs up to 2 m (much smaller at higher altitudes). Stem and leaves covered with stellate hairs often turning fulvous when dry. Leaves alternate, short-petiolate, entire, linear, oblong or ovate, basally cuneate to slightly cordate, green. Flowers unisexual (plants monoecious); male flowers agglomerated in dense spike-like inflorescences that terminate the branches, perianth hyaline with 3-5 almost free segments covered with stellate but easily caducous indumentum, stamens 3-5; female flowers positioned below the male ones, enclosed by a cover consisting of two at least halfway fused accrescent bracts with long simple hairs and much smaller stellate hairs. Styles 2. Fruits with hyaline, very thin pericarp covered with scattered stellate hairs. Seeds ~2.0 mm with thin seed coat, perisperm abundant. Embryo vertical, horseshoe-shaped.

Axyris villosa
One species in the deserts of Eurasia and North America. According to recent investigations (Heklau and Röser 2008), the genus comprises one species. Herb. Linn. 1101.1 (LINN! image available at http://linnean-online.org/11529/). The full synonymy is given by Heklau and Röser (2008).
Taxonomic notes. Morphologically, two forms of K. ceratoides can be distinguished based on the shape of the leaf base; some specimens in Himalaya have slightly cordate or truncate leaves while others have cuneate leaves. In Tibet, a form with cuneate leaves is found. Description. Glabrous or scarcely pubescent annual up to 50(70) cm, branches often terminating in acicular apices. Leaves long-petiolate, up to 10.0 cm, broadly ovate or ovoid, dentate or erose, teeth straight or incurved, tip mucronate, blades glabrous or sparsely covered with simple, often curved hairs. Inflorescences in the leaf axils, quite short, falsely dichotomous. Perianth of 5 free segments, keeled along midrib. Stamens 2, anthers small, 0.2-0.3 mm, without appendages. Stylodia concrescent into column in their lower half. Fruit dehiscent by a lid; pericarp white or greenish with several homocellular layers. Seeds dark-red, depressed-globular, ~1.3 mm in diameter, smooth.
One species in Himalaya and West and Central China.

Note.
Although Amaranthus diandrus has a very clear diagnosis (Sprengel 1822), it was an unresolved name (Das 2016) and is now added to the synonyms of Acroglochin persicarioides.
Description. See the genus description.
Description. Annuals, branched from the base, often forming tumble-weed habit, covered with branched hairs, usually glabrescent at the fruiting stage. Leaves sessile, filiform to ovoid, green or greyish, sometimes red from both sides. Perianth absent or of 1-3 hyaline segments. Stamens 1-3, anthers 0.2-0.3 mm long. Stylodia 2 with persistent lower part. Fruit ovoid or roundish, 1.5-6.5 mm, glabrous or covered with branched (usually caducous) hairs, adaxially plain or slightly concave, abaxially convex; marginal wing membranous, well-developed or tiny. Fruit apex roundish, triangular or emarginate. Pericarp either adherent to the seed coat or its outer layer forms wart-like or (rarely) wavy detachments; groups of dark-brown cells filled with tannins are often present in the outer pericarp layer. Seed coat of two or more thin layers.
Fruit anatomy plays the most important role in the diagnostics of the species groups. The following characters can be noted (Sukhorukov 2007b): (1) thickness of the fruit, (2) thickness of the outer pericarp layer, (3) presence of sclereids in the medial fruit part and their quantity and orientation and (4) outline and size of the fruit wing.
The exact species number is still unknown, but it is estimated at approximately 80 (Sukhorukov 2014).
All species encountered in our area are alpine plants growing at elevations of 2800-5000 m.   Iljin (1929) indicates two type specimens ("Kew [K] and Vienna Univ. [WU]") in the protologue. We did not find the material in WU and thus select a lectotype in herbarium K with additional authentic material in other herbaria.
Description. Annual up to 40 cm, slightly pubescent. Leaves lanceolate, oblanceolate or narrowly oblong, often recurved, up to 3.0 × 0.5 cm. Inflorescence elongated, with many flowers. Bracts usually recurved, sickle-shaped, not completely covering the fruit. Fruits 3.5-4.0 × 3.0 mm, glabrous but sometimes with dark-brown cells and scattered warts (Fig. 33A); fruit margin emarginate at the top. Wing 0.45-0.60 mm, denticulate or crisp, narrowly triangular in cross-section. Sclereids in the medium fruit part consist of 1-2 cell layers orientated parallel to the fruit axis.
Habitat Description. Dwarfish, densely pubescent annual up to 6 cm with a single stem or with short lateral branches. Leaves linear, up to 1.5(2.0) mm wide. Inflorescence very short, with several flowers. Bracts slightly shorter than fruits (fruit margins visible). Fruit ovoid, 2.5-2.7 × 2.0 mm, glabrous, apex rounded and only slightly emarginated General distribution. Endemic to South Tibet. Description. Annual up to 20 cm, usually pubescent. Leaves linear to narrowly oblong, up to 2.0 × 0.5(0.7) cm, with a mucro at the top. Inflorescence short, its branches with 5-15 flowers. Bracts significantly shorter than leaves, ovoid, (almost) completely covering the fruit, sometimes recurved at the fruiting stage. Fruit 2.2-3.0 × 1.7-2.0 mm, 0.40-0.60 mm thick, with entire margin and indistinctly triangular or roundish apex terminating in slightly emarginate beak (Fig. 33C), glabrous and mostly smooth (without detachments), sometimes with groups of dark-brown tanniniferous cells. Wing small, 0.15-0.30 mm, broadly triangular in cross-section. Pericarp without detachments. Sclereids in the medium fruit part composed of 1-2 layers located parallel to the fruit axis.
Description. Annual up to 20 cm, glabrescent with ascending branches. Leaves linear to oblong (the lower leaves often spatulate), up to 3.0 × 0.6 cm, often recurved with visible white margins and a mucro at the top. Inflorescence elongated, with many flowers. Bracts leaf-like, lanceolate, significantly exceeding the fruit in length and covering it completely laterally. Fruit 2.8-3.5(4.0) × 2.0-2.5 mm, flat or often folded, acute (triangular) apex with verrucose (warty) surface, without stellate hairs (type variety) or with stellate hairs (var. montanum) and clearly visible (0.45-0.70 mm) undulate wing that is narrowly triangular in cross-section. Pericarp detachments up to 100 µm. Sclereids in medium fruit part with 1-2 layers located parallel to the fruit axis, rarely with no sclereids.
Distribution. Corispermum dutreuilii seems to be the most common species of the genus in Ladakh and NW Xizang. Both varieties can grow together, e.g. in Ladakh (collection numbers 3465 & 3470 for the type variety and 6265 & 6266 for var. montanum (PRA)).

Corispermum pseudofalcatum
Habitat. Screes and sandy riverbeds; 3000-4200 m a.s.l. Phenology. Flowering: July-September; fruiting: August-October. Distribution. This species is known only from the type locality (Fig. 35). Both morphological and carpological characters as well as its record in South Tibet indicate its affinity to C. lhasaense.  Description. Annual up to 20 cm, glabrescent. Leaves lanceolate, up to 3.0 × 0.3 cm long. Inflorescence dense, 3-5 cm long. Inflorescence elongated. Bracts oblong to ovoid, moderately recurved or sometimes appressed to the stem, completely cov-
Description. Annual up to 20 cm, glabrescent. Leaves linear to narrowly oblong, up to 3.0 × 0.5(0.7) cm. Inflorescence short. Bracts shorter than leaves, lanceolate, (almost) completely covering the fruit, appressed to the stem. Fruit 2.6-3.5 × 1.7-2.0 mm, 0.40-0.60 mm thick, flat or folded adaxially, with triangular apex terminating in slightly emarginate beak, with stellate hairs and mostly warty (Fig. 36C), sometimes with groups of dark-brown tanniniferous cells. Wing 0.15-0.45 mm, entire to dentate, broadly triangular in cross-section. Sclereids in the medium fruit part composed of 1-2 layers located parallel to the fruit axis.
Habitat. Sandy places at altitudes of 3500-4800 m. Corispermum tibeticum Iljin, Izv. Glavn. Bot. Sada SSSR 28: 644 (1929). Lectotype (Sukhorukov, designated  In the protologue (Iljin 1929), the specimens were cited as a type material of C. tibeticum, but the Falconer syntype kept at K is not informative and rather deviates from the typical C. tibeticum by its almost wingless fruits. We propose to select a lectotype from other specimens (syntypes) cited by Iljin in the protologue.  there is a significant gap between the distribution of typically alpine A. tibeticum and all other species of the genus.
Description. Annual up to 20(30) cm with tumble-weed habit; stem very densely pubescent with dendroid hairs (up to 2.5 mm in length, Fig. 39A) and often whitish when young with partially persistent hairs in the fruiting stage. Lower leaves oblong or oblong-lanceolate, continuously decreasing in size and shape in the inflorescence, 2.0-8.0 × 0.5-1.5 cm, tapered in petiole, with (5)6-10 prominent nerves, densely pubescent when young. Inflorescence leafy; flower clusters remote in the lower part of the inflorescence and densely arranged in the upper part. Anthers ~0.3 mm. Pericarp covered with dendroid hairs, glabrescent at fruiting. Seed lens-shaped (Fig. 39B, C), 1.9-2.1 mm × 1.3-1.4 mm, yellow with brownish tanniniferous spots.
Differs from the two related species, A. pungens (Vahl) Link ex A.Dietr. and A. gobicum Bunge (if the latter is accepted at the specific rank), by persistent dendroid pubescence (with longer hairs) in the inflorescence at the fruiting stage as well as by its disjunct distribution in South Tibet.
Habitat. Sand dunes and stony riverbeds; 2800-3700 m a.s.l. Phenology. Flowering: August-September; fruiting: September-early October.  Taxonomic notes. The examined specimens have perianth with unequal projections at the fruiting stage that is consistent with the species description by Paulsen (1903). However, Lomonosova and Freitag (2008)   Description. Annual up to 25 cm with often reddish stems covered with spreading hairs. Leaves up to 1.0 cm, oblong to ovate, densely covered with (semi)appressed hairs (both Bassia scoparia and other Chenopodiaceae. However, the presence of B. indica is possible in the foothills of North Himalaya (0-1500 m a.s.l.), as it is known from neighbouring areas, e.g. state of Haryana, India (BSD! LE!) and Punjab Province, Pakistan (K!). Description. Annuals, usually with tumble-weed habit. Stems and leaves with white, simple multicellular hairs that are appressed and obliquely orientated and with scattered yellow short hairs mostly located in the leaf axils. Leaves terete, green, obtuse, with water-storage tissue and C 3 -type anatomy. Flowers in leaf axils, solitary or in pairs. Perianth of 5 at least half-way concrescent segments, usually pilose; all or three of five segments develop wing-like or spiny projections. Stamens 5, with small anthers. Fruit with hyaline pericarp adherent to the seed coat. Seed embryo horizontal. Perisperm abundant.

Grubovia
Here, we indicate for the first time that the presence of short yellow hairs is the remarkable morphological characteristic of Grubovia that is important for the delimitation of this genus. Grubovia comprises three species mostly distributed in Central Asia. Grubovia dasyphylla has the largest distribution that also includes Tibet and North Himalaya.  Taxonomic notes. The taxa related to Salsola kali were considered within the reinstated genus Kali (Akhani et al. 2007), which was accepted in subsequent taxonomic treatments (Gutermann 2011, Sukhorukov 2014, Sukhorukov et al. 2016b). However, Mosyakin et al. (2014Mosyakin et al. ( , 2017 presented some arguments in favour of a recent nomenclatural proposal to conserve the generic name Salsola with S. kali (=Kali turgidum) as a conserved type. The Nomenclature Committee for Vascular Plants did not reach a consensus regarding the recommenda-tion or rejection of this proposal (Applequist 2016), but the accepted type of the genus Salsola is S. kali under the new Art. 10.5-10.7 of the ICN (Turland et al. 2018). Description. Annuals or subshrubs, glabrous or papillate, with tufts of simple curved hairs in the leaf axils. Leaves mostly alternate or lower leaves opposite, semi-terete or terete, sometimes flattened, usually less than 5 mm in width, stiff or fleshy, with a persistent yellowish mucro up to 3.5-4.0 mm long. Bracts longer than bracteoles or equal in size. Flowers of two types: some located below the main inflorescence and arranged in clusters mostly consisting of two female flowers supported by concrescent and basally hardened bracts and bracteoles, rarely clusters one-flowered (these fall off with bracts and bracteoles), perianth r-shaped, usually with small wings, tubercles or without any projections in the flexure; flowers of the second type grouped in the main inflorescence, hermaphrodite, each flower supported by a free bract and two bracteoles (rarely concrescent), r-shaped or tubuliform at fruiting (in the latter case, the segments in their upper part are convergent, forming a stout or hyaline conus that covers the fruit from above). Perianth of (4)5 segments, with well-developed, equal or unequal wings or tubercles at the fruiting stage. Stamens (in the hermaphrodite flowers) 5, divided in the lower half, without prominent appendages at the tip of the anthers. Stigmas 2, usually equal to style (or a style is very short). Fruits dry or somewhat fleshy in the upper part. Seeds with horizontal or obliquely orientated embryo (sometimes seems to be vertical due to anacrostyly). Perisperm absent.
Approximately 22 species in the steppes, seashores and deserts of Eurasia, North America, Africa and Australia.
Key to the species 1 Bracts appressed to the stem; perianth (of the flowers located in the main inflorescence) at fruiting with tubercles; alternatively, two (out of five) perianth segments bear short wings up to 1. Taxonomic notes. The new species was formerly identified as S. jacquemontii or S. collina. In many characters (e.g. tiny anthers, small outgrowths of the perianth segments), Salsola hartmannii resembles S. monoptera, but the latter species has only one well-expressed, wing-like projection on a segment closest to the bract, whereas projections on other segments are much smaller (in the form of tubercles). Additionally, the gibbous bracts and bracteoles are exceptionally rare in S. monoptera. In contrast, S. hartmannii possesses gibbous bracts and bracteoles and two or three perianth segments develop tiny, but clearly visible wings. The new species is also similar to S. jacquemontii but has smaller anthers and a smaller diameter of the entire diaspore (winged perianth enclosing the fruit), which is hidden within the subtending bract and bracteoles.
The gibbous bract and bracteoles cannot be considered as strong characteristic features of S. hartmannii; some specimens of S. jacquemontii may possess such a shape of the bracts and bracteoles. All three species, S. monoptera, S. hartmannii and S. jacquemontii, are morphologically related. Salsola collina is morphologically distant from these species, having appressed bracts without clearly expressed wings on the perianth at fruiting. The distribution patterns are also different: S. monoptera is widespread in Central Asia and Tibet, S. hartmannii seems to be localised in North Himalaya and Karakoram and S. jacquemontii is found in Central Himalaya and Tibet. The presence of S. jacquemontii and S. collina in North Himalaya (Rilke 1999) is not confirmed.
Description. Annual with prostrate or ascending stems up to 15 cm long, glabrous or scarcely papillate. Leaves terete, 10-20 × 1.0-1.5 mm. Flowers in axillary clusters and arranged in the main inflorescence. Flower clusters (located below the main inflorescence) abundant and consist of two flowers covered with bracts and bracteoles that are connate almost to the top and basally gibbous; perianth segments membranous, fimbriate, r-shaped, their tips more or less appressed to each other, not hardened and not forming a tight conus, with white, small tubercles 0.3-0.5 mm long in the flexure; styles with stigmas ca. 0.5 mm; fruits 1.2-1.6 mm, depressed roundish. Flowers in the main inflorescence with (almost) free, basally gibbous bract and bracteoles, hermaphrodite; perianth segments 5, membranous, fimbriate, r-shaped at the fruiting stage, < 2.5 mm long, above the flexure soft, hyaline, horizontally ori- entated and not forming a distinct conus; 2 or 3 (of 5) segments develop wing-like appendages (up to 1.5 mm long) that are located almost horizontally or obliquely, two other segments develop tubercles only; the whole diaspore with the perianth wings is 2.0-3.0 mm across and is hidden within the bract and bracteoles; anthers 0.3-0.5 mm long; styles with stigmas ca. 1 mm; fruits lenticular, 1.1-1.5 mm. Seeds with horizontal or obliquely orientated embryo.
Habitat. Gravelly or sandy substrates at altitudes of 3500-5200 m. The species seems to be common in Ladakh and Spiti Valley.  General distribution. North India, NW China (Xizang) and North Pakistan ([Gilgit-Baltistan] Nagar, dry stony slope, 9000 ft a.s.l., 22 Aug 1987, Jamshed 239 (E!)). Description. Erect or prostrate (in mountainous areas) annual up to 70 cm, stems reddish, glabrous or scarcely papillate. Lower leaves up to 80.0 × 0.5-1.0 mm, much shorter in the middle and upper stem parts, linear or filiform, with a mucro 0.6-2.0 mm long. Flowers in axillary clusters and arranged in the main inflorescence. Flower clusters (located below the main inflorescence) abundant, consisting of two flowers surrounded by bracts and bracteoles that are connate almost to the top and not gibbous basally; perianth segments of each flower membranous, glabrous, r-shaped, their tips appressed to each other, hardened and not forming a tight conus, with white, small tubercles 0.3-0.5 mm long or without any projections in the flexure; styles with stigmas ca. 1.0 mm; fruits 1.0-1.3 mm, depressed roundish. Flowers in the main inflorescence with (almost) free, basally not gibbous bract and bracteoles (that are appressed to the stem), hermaphrodite; perianth segments membranous, glabrous, r-shaped at fruiting stage, (1.5)2.0-2.5 mm long, membranous above the flexure and not forming a distinct conus; two abaxial segments usually have short erose wings (up to 1.0(1.5) × 1.5 mm), their tips (above outgrowths) hyaline and not stout; anthers 0.6-0.9 mm long; styles with stigmas ca. 1.0 mm; fruits lenticular, 1.1-1.5 mm. Seed seems to be vertical (due to anacrostyly) but is indeed horizontal. inflorescence. Clusters (located below the main inflorescence) consist of two female flowers supported by a bract and two bracteoles that are connate almost to the top and usually not gibbous at the base; perianth segments membranous, fimbriate, r-shaped with their tips appressed to each other, not hardened and not forming a tight conus, with small, reddish or white tubercles up to 0.6 mm long in the flexure; styles with stigmas ca. 1.0 mm long; fruits 1.3-1.6 mm in diameter. Flowers in the main inflorescence with (almost) free, usually not gibbous, bract and bracteoles, hermaphrodite; perianth segments membranous, fimbriate, r-shaped at fruiting stage, < 2.5 mm long, soft above the flexure, hyaline, 0.6-0.8 mm long, horizontally appressed and not forming a distinct conus; three segments with horizontally orientated wing-like appendages (< 3.0 mm long), two other segments with tubercles only (the entire diaspore with the perianth wings is 3.5-6.0 mm across); anthers 0.5-0.7 mm long; styles with stigmas ca. 1.0-1.5 mm long; fruits lenticular, 1.1-1.5 mm in diameter. Seeds with horizontal or obliquely orientated embryo.

Salsola collina
Habitat. Gravelly substrates, limestone at elevations of 2600-4200 m. Phenology. Flowering: July-September; fruiting: August-October.   3 (LINN). The list of the synonyms is extremely large for this species (Rilke 1999), but some varieties and species described from Europe or Central Asia require additional study.
Description. Annual up to 70 cm high. Stem branched from the base, finely papillate or glabrous as well as the leaves. Leaves 20.0-80.0 × 1.0-2.0 mm, linear, with a mucro (1.5)2.0-3.5 mm long. Flowers in axillary clusters and arranged in the main inflorescence. Clusters (usually located below the main inflorescence) consist of two often unequal female flowers, each flower supported by a bract and two bracteoles and additional bracts are sometimes present in the clusters; bracts and bracteoles fused in their basal part; perianth segments free, membranous, r-shaped, their tips appressed to each other, not hardened, not forming a conus, with small or hardly noticeable, red to pinkish wings in the flexure; styles with stigmas ca. 1.5 mm long; fruits 1.0-1.6 mm, depressed roundish to ovoid. Flowers in the main inflorescence with a free bract and two bracteoles (shorter than bract or equal in length), hermaphrodite; perianth segments membranous, tubuliform (not r-shaped), 3.0-4.0 mm long at fruiting stage, their lower part marginally glabrous or minutely fimbriate; segment tips hyaline, forming a convergent, glabrous conus 1.0-1.5 mm long with well-developed, horizontally located or slightly recurved, entire to crisp wings in the flexure; three outer segments develop roundish wings and two inner segments have narrower, lanceolate outgrowths (the whole diaspore with the perianth wings is ( Taxonomic notes. The hardness of the conus (perianth part above the wings or tubercles at the fruiting stage) is a very important trait for identifying the representatives of the Salsola type section (ex-Kali), which is widely used in taxonomic treatments (e.g. Iljin 1936, Tzvelev 1996, Rilke 1999. The length of the conus seems to be another important characteristic for the taxonomy of this group. All specimens of S. austrotibetica were formerly identified as Salsola ikonnikovii or rarely S. tragus (incl. S. pestifer). The identification of the Tibetan specimens as S. ikonnikovii (Rilke 1999) was mostly based on the similar broader and stiff leaves. Salsola ikonnikovii was formerly considered more broadly with a vast distribution covering Central Asia and Tibet (Rilke 1999). Although some other Central Asiatic species were included in S. ikonnikovii (S. beticolor Iljin, S. centralasiatica Iljin, S. microkali Popov, S. potaninii Iljin and S. chinghaiensis A.J.Li), the specific status of these taxa must be re-evaluated following extensive molecular and morphological investigations. All southernmost records of Salsola ikonnikovii cited by Rilke (1999) from Tibet [Xizang] are here considered Salsola austrotibetica. Salsola ikonnikovii s. str. is characterised by a large (10-20 cm) main inflorescence with very densely arranged flowers, winged perianth 5.0-8.0 mm in diameter and peri- anth segments forming a narrowly triangular and stiff conus of (2)2.5-3.5 mm long. In contrast, the new species has a much smaller (up to 12 cm) inflorescence, perianth with wings 4.0-5.0 mm across and triangular and slightly hardened perianth conus 1.5-2.0 mm long. All plants in the PE herbarium were identified as Salsola ruthenica [≡S. tragus], another similar-looking species. However, linear or filiform leaves and a completely hyaline perianth conus distinguish S. tragus from both S. ikonnikovii and S. austrotibetica.
The distribution area of Salsola austrotibetica is restricted to South Tibet (Fig. 48) and that of S. ikonnikovii covers Mongolia and the northernmost provinces of China (Nei Mongol, Ningxia and Gansu).
Description. Annual up to 40 cm high. Stem straight, branched from the base with ascending lateral shoots; stem and leaves finely papillate. Leaves on the main stem remote (with internodes 20-40 mm long), 20-50 mm long, 1.0-4.0(5.0) mm wide above the broad base, slightly flattened, lanceolate to linear, thick, with a mucro (1.0)1.5-2.5 mm long. Flowers in axillary clusters and arranged in the main inflorescence. Flower clusters (usually located below the main inflorescence) consist of two often unequal female flowers, each flower supported by a bract and two bracteoles, sometimes additional bracts are present in the clusters; bracts and bracteoles fused in their basal part; perianth segments of each flower membranous, r-shaped, their tips appressed to each other, not hardened, not forming a conus, with red to pinkish, small or hardly noticeable wing-like tubercles in the flexure; styles with stigmas ca. 1.5 mm long; fruits 1.0 to 1.6 mm, depressed roundish to ovoid. Flowers in the main inflorescence lax, hermaphrodite, with two bracteoles (slightly shorter than bract) and a free bract having a broad (up to 1.0 mm) white margin; perianth segments membranous, tubuliform (not r-shaped), 1.7-2.2 mm at flowering stage and elongated to 4.0-5.0 mm at fruiting, their lower part marginally glabrous or finely fimbriate; segment tips slightly hardened but not hyaline, forming a convergent, glabrous conus 1.5-2.0 mm long, with well-developed, obliquely orientated, unequal, erose to lobate wings in the flexure; the whole diaspore with the perianth wings is 4.0-5.0 mm across; anthers General distribution. Central Asia and South Siberia. The species was probably first collected in Xizang in the 1980s-1990s (Rilke 1999), but it is not mentioned in Zhu et al. (2003). The records cited below expand the distribution of S. monoptera to the south (compare to Rilke 1999). Description. Annual up to 30 cm with a tumble-weed habit at the fruiting stage; young plants often woolly due to abundant curved simple hairs in the leaf axils. Stem and leaves glabrous or shortly papillate. Leaves terete, up to 15 mm long, with a yellowish, easily caducous mucro up to 5.0 mm. Flowers covered with a bract and two bracteoles with two types of perianth (most of the flowers with five hyaline perianth segments (up to 3.0 mm long) with an orbicular white or pink wing (2.0-3.0 mm, sometimes 0.7-1.5 mm in diameter) near their tips; a small part of the flowers with 5 perianth segments hardened almost to the top without any projections or tuberculate). Stamens 2-3. Anthers 0.5-0.6 mm long. Fruit 1.0-1.2 mm. Seeds with vertical embryo, with the radicula pointing upwards. Perisperm absent.
Description. Sub-shrublet up to 30 cm with a robust caudex, branched from the base. Stems glabrous or very shortly papillate, with two furrows. Lower leaves up to 30-35 mm long, fleshy, semi-terete, appressed to the stem or rarely recurved with tufts of hairs in their axils, without mucro or with a hardly noticeable mucro at the top. Bracts usually longer or equal to the flowers. Flowers solitary in the axil of the bract. Perianth segments greenish, 5, at fruiting stage with wings 2.0-2.5 × 2.5-3.5 mm, purple or lilac turning brownish. Anthers 5, ~1 mm long, ovate, apically without an appendage. Style short, 0.25 mm with 2-3 stigmas of the same length. Fruit whitish, depressed orbicular, ~1.5 mm in diameter.
Habitat. Rocky areas, semi-deserts and deserts at high elevations (3500-5200 m). Phenology. Flowering: July-September; fruiting: September-October. Distribution. See Fig. 53. easily distinguished by the following leaf characteristics: the leaves of all Halocharis (tribe Caroxyleae) are covered with both short and long simple hairs, while those of Halogeton (tribe Salsoleae) are glabrous or covered with short papillae (long, clearly visible hairs are present only at the leaf base). The presence of two hair types is a distinct characteristic of Halocharis compared with many other Salsoloideae (Sukhorukov et al. 2016b).