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Elwendia is a middle-sized genus with its center of diversity in Central Asia. Due to the progress in recent molecular phylogenetic studies, the genus has been restored and expanded by the inclusion of eastern species of Bunium. However, it still lacks a modern infrageneric classification system based on integrative analyses of molecular, morphological, and karyological evidence. Here, we present the most comprehensive phylogenetic analysis of the genus to date, covering all species within the genus. We used DNA sequences from the nuclear ribosomal internal and external transcribed spacers and sampled 21 morphological characters for each species. As a result, an infrageneric classification system of Elwendia is proposed with 6 sections and 11 subsections. Diagnostically important morphological characters include specializations of fruits and stem leaves. Of particular interest is the congruence between molecular and chromosomal data. Descriptions and species composition of the sections and subsections are provided. Elwendia longiloba is resurrected from the synonymy of E. hissarica.

Maximum Likelihood (ML) and Bayesian analyses (BI) applied for combined plastid matrix (cpDNA trnL(UAA) intron, trnL–trnF IGS, and rpl32–trnL(UAG) IGS regions) of 124 accessions of 36 species of the genus Atraphaxis have shown new species from southern Siberia. The analysis of 112 accessions of 36 species of nrDNA ITS1–5.8S–ITS2 regions showed less clear topology. A new species A. kuvaevii endemic for Yenisey Valley in Tuva was revealed with molecular and morphological data. The plants attributed previously to A. pungens split into three cryptic species with different plastid regions. A. pungens grows in the south of Krasnoyarskii Krai, Khakassia, Tuva and Mongolia. A. ledebourii is distributed in Altai Mts, Tuva and Mongolia. It combines a functional ITS ribotype and a psedogenized ribotype, what agrees with high chromosome numbers. The third pseudogenized ribotype shared with A. ariana was found in a single sample from SE Tuva. A. ×uyukensis is a species of hybrid origin from the Uyuk ridge with the ITS ribotype of A. pungens, which probably captured cpDNA of unknown species related to A. selengensis and A. davurica by maternal lineage. Despite the differences in plastid regions, these cryptic species have no clear morphoplogical distinctions. The hybrid A. pungens × A. ledebourii from Tuva and the hybrid A. ledebourii × A. pungens from Hangay Mts in Mongolia do not differ from the parents. The eastern race of A. frutescens s.l. sub A. decipiens from Tuva and Altai Republics differs from the western race of A. frutescens with a single nucleotide substitution in cpDNA trnL–trnF IGS, a distinct ITS ribotype, a short perianth tube equal to outer tepals, herbaceous tubular ochreolas and linear-lanceolate leaf blades. The hybrids A. pungens × A. decipiens and A. decipiens × A. pungens were detected in three locations in Tuva. Cryptic hybrids A. ledebourii × A. decipiens, A. ledebourii × A. frutescens, and A. decipiens × A. ledebourii were detected in Altai and Tuva Republics. Two hybrids A. decipiens × A. ledebourii have shown the capture of cpDNA of A. decipiens. Atraphaxis laetevirens is limited by the territory of Kazakhstan, Altai Republic and Dzungaria and does not grow in Tuva and Khakassia.

Elwendia Boissier (1844: 140). Type:— Elwendia caroides Boissier (1844: 140). Corrected description:—Perennial polycarpic, rarely monocarpic herbs. Tubers not branched, globose or ovate, deeply submerged in soil. Stems solitary, solid or hollow, glabrous, branched in upper part. Leaves twice or three times dissected. Sheaths of stem leaves usually not inflated. Petioles glabrous. Rachis not refracted or refracted. Primary segments of leaves petiolulate. Terminal leaflets filiform, lanceolate to ovate. Terminal umbels broader than lateral, without involucre, or more rarely with a few bracts. Involucel of entire, filiform to linear, herbaceous or white-margined bracteoles. Pedicels during fructification thin ( 0.7 mm in diam.), very short (up to 3 mm) to long (11–20 mm). Calyx teeth absent. Petals white. Fruits glabrous, separating into, usually from below, rarely (E. chaerophylloides) from above, mericarps or, rarely not separating into mericarps. Mericarps orbicular, elliptic to linear, with or without beak, slightly compressed or convex dorsally. Dorsal mericarp ribs almost obsolete to shortwinged. Marginal mericarp ribs usually not winged, rarely winged. Stylopods short conical or conical. Styles recurved, spreading or erect. Exocarp of small cells. Commissure rather broad (exocarp interrupts near edges of marginal ribs) or intermediate (exocarp interrupts in middle part of commissure). Vallecular vittae usually solitary, rarer 2–3 per furrow or numerous. Endosperm on commissural side usually flat, rarely (E. angreni, E. badachschanica, E. setacea, E. stewartiana, E. vaginata) grooved. Embryo and seedlings with two normally developed cotyledons. Geographic distribution:—From Central Iran to Western China, Mongolia, and the Himalayas, with the center of diversity in the Pamir-Alay. Species number:—30., Published as part of Kljuykov, Eugene V., Zakharova, Ekaterina A., Degtjareva, Galina V. & Samigullin, Tahir H., 2023, Infrageneric classification of Elwendia (Apiaceae) with a restored species, pp. 149-168 in Phytotaxa 595 (2) on page 157, DOI: 10.11646/phytotaxa.595.2.3, http://zenodo.org/record/7905847, {"references":["Boissier, E. (1844) Plantae Aucherianae. Adjunctis nonnullis e regionibus Mediterraneis et Orientalibus aliis cum novarum specierum descriptione. Annales des Sciences Naturelles. Botanique, ser. 3, 1: 120 - 151."]}

1. Elwendia sect. Elwendia. Type:— Elwendia caroides Boissier (1844: 140). Description:—Polycarpic herbs; stems solid; leaf rachis and petiolules not refracted; stem leaf sheaths not inflated; upper stem leaves pinnate with straight rachis; umbels corymbose or hemispherical, without involucre; bracteoles present; pedicels during fructification thick; fruits hardly or rarely easily separated into mericarps; dorsal mericarp ribs keeled; marginal mericarp ribs winged or not winged; mericarps with a beak under stylopodia (Fig. 2A); styles recurved or erect; commissure broad, exocarp terminates near edges of marginal ribs (Fig. 3A); mesocarp in marginal ribs consists of non-lignified parenchyma cells or lignified parenchyma cells with pitted walls; vallecular vittae solitary, 2–3 per furrow or numerous; endosperm on commissural side flat; 2n = 12, 13, 22. Note:—The delimitation of the section follows the original sectional concept of Kljuykov (1988). The diagnostic feature of this section is a presence of the beak under stylopodia (Fig. 2A), which is distinct from other Elwendia species., Published as part of Kljuykov, Eugene V., Zakharova, Ekaterina A., Degtjareva, Galina V. & Samigullin, Tahir H., 2023, Infrageneric classification of Elwendia (Apiaceae) with a restored species, pp. 149-168 in Phytotaxa 595 (2) on pages 157-158, DOI: 10.11646/phytotaxa.595.2.3, http://zenodo.org/record/7905847, {"references":["Boissier, E. (1844) Plantae Aucherianae. Adjunctis nonnullis e regionibus Mediterraneis et Orientalibus aliis cum novarum specierum descriptione. Annales des Sciences Naturelles. Botanique, ser. 3, 1: 120 - 151.","Kljuykov, E. V. (1988) A survey of the genus Bunium L. Revision of the generic system. Byulleten' Moskovskogo Obshchestva Ispytatelei Prirody. Otdel biologicheskii 93: 76 - 89. [In Russian]"]}

Aulacospermum multicaule Pimenov & Tojibaev, sp. nov. (Figs. 1, 2A, 3, 4A, 5, 7) Type:— UZBEKISTAN. Western Tian-Shan, Kurama Range, Chadaksay River Basin, upper reaches of Pashakhansay, locality Betagadik, slopes to the say, between rocks, on fine earth, 07 July 2018, K .Tojibaev, N. Beshko & D.Makhmutzhanov s.n. (holotype TASH, isotype MW). Aulacospermum multicaule is similar to A. roseum Korovin but differs from the latter in numerous (up to 11) branched stems, mostly black remains of petioles and sheaths of leaves of the preceding year at rootstock, and petal colour (yellowish vs. white). Species nostra Aulacospermo roseo Korovin similis, sed plantis caespitosis, caulibus numerosis (usque ad 11), ramosis, reliquiis petiolorum vaginisque foliorum anni praecedentis caudicibus obtectis fere nigris, squamiformibus, membranaceis (non fibrillosis), petalis flavescentibus (non albis) differt. Erect, multi-stemmed (up to 9‒11), polycarpic perennial. Rootstock branched, with dark brown or almost black, membranous, non-fibrillating remains of last year’s petioles and sheaths of basal leaves. Taproots vertical slightly branched. Stems 18‒42 cm tall, solid, finely grooved, twig-like, 3‒5 mm diameter at base, glabrous, branched almost from the base, with white parenchymal stem core. Leaves predominantly basal, forming dense rosettes or located in the very lower part of stems, with short petioles slightly inflated into narrow sheaths. Leaf blades oblong in outline, 5‒7 cm long, doubly pinnatisect with 2–4 pairs of lateral segments; terminal leaflets lanceolate, with cuneate base, 5‒7 mm long, slightly pointed. Cauline leaves small, glabrous, entire or ternate, sessile, with short narrow sheaths. Petioles glabrous, terete or almost adaxially flattened, with a broad shallow adaxial groove, a small central cavity, with 5 peripheral vascular bundles, 3 peripheral collenchyma bundles and a small collenchyma bundle along the adaxial groove. Umbels sparse, 2‒4.5 cm diameter, 3‒5(6)-rayed, rays unequal, 2‒5(6) cm long, thin, curved. Bracts 1‒3, unequal, linear or linear-lanceolate, early falling. Umbellules compact, 0.5‒1 cm diameter, 3‒5-flowered. Bracteoles 2‒3, short triangular-lanceolate, membranous, with narrow white margin. Pedicels short, nearly equal. Petals yellowish. Mericarps homomorphic, elliptic or ovoid in outline, 4‒5 mm long, 2.5‒3 mm wide, only primary ribs present. Mericarp surface with scattered multicellular tubercles, colliculose, carpophore bifurcated to base; mericarp ribs equal, winged, straight or slightly undulate. Calyx teeth conspicuous, broadly triangular, remaining at fruit, with white membranous margin. Stylopodium low-conical or flat. Styles recurved. Exocarp cells large (40‒50 μm), with convex outer walls. Commissure narrow, exocarp almost reaching carpophore. Mesocarp parenchyma not lignified; vittae thin and solitary in valleculae, two on commissure; rib secretory ducts absent; endosperm on commissural side with a narrow deep groove. Etymology:— The species epithet refers to numerous branched stems. Phenology:— The plant was collected with unripe fruits at the beginning of July; most probably, flowering in June, fruiting in July. Distribution and ecology:— Aulacospermum multicaule is most probably endemic to Uzbekistan (Fig. 8). It grows between rocks on fine earth.

Atraphaxis pungens, A. ledebourii sp. nov. and A. × uyukensis nom. provis. Our current results have shown that “ Atraphaxis pungens s.l. ” combines plants from Southern Siberia and Mongolia with three different plastid haplotypes, which replace each other in the area (Figs 2 and 8). All the plants demonstrate a great similarity in the appearance and morphology of shoots, inflorescences, perianths and fruits, so it is impossible to discriminate them. They are shrubs 15–50 cm high (Figs 9–11, Table 1) with vigorous vegetative shoots, often sylleptically branching. Second-year shoots become durable, long lasting, perennial axes bear prickly branchlets. The covering of second-year shoots is dark-gray, almost black, the cork peels off, exposing the reddish bark. Wood is pale pinkish. Second-year shoots and their branchlets bear short (5–10 cm long) axillary pale-green or creamy annual shoots. These shoots are leafy and some terminate with thyrses. Annual short generative and vegetative shoots bear small oblanceolate or spathulate leaf blades with acute or obtuse apex, gradually narrowed to the base (Fig. 9A–C; Fig. 10A, C, D; Fig. 11; Fig. 12A–B; Fig. 13A, Fig. 14D, E). Annual long vegetative shoots bear large obovate, oblongelliptic, broadly elliptic or rhomboid-elliptic leaf blades with acute or mucronate apex (Fig. 9D–F; Fig. 10B, E, F; Fig. 12C; Fig. 13B; Fig. 14A). The thyrses are usually compact, frondo-bracteose, consist of 4–7 cymes with 2–3 flowers. Perianth is yellowish-white or white, sometimes pinkish, with a very long perianth tube articulated to as long as pedicel (Fig. 12H, I; Fig. 13C; Fig. 14B, F). The epidermal cells of tepals and tube are similar (Fig. 6C–F). Atraphaxis pungens has black glossy fruits twice shorter than inner tepals of the perianth (Fig. 12J–L). Atraphaxis ledebourii has dark-brown to black fruits, 1.5 times shorter of inner tepals (Fig. 13F, G, I). In plants from Altai and Tuva, the ovary and stamens are often underdeveloped. In both species three styles are fused at the very base (Fig. 6J, K, Fig. 12D–G, Fig. 13D, E, H). In A. pungens and A. × uyukensis, stigmas are either flat and rounded, or elongated and bent out, later bevealed outward (Fig. 12G, L; Fig. 14C, G–I). In A. ledebourii from Altai Republic, the stygmas are flat and rounded (Fig. 13D), in plants from Tuva Basin and Ubsunur Basin the stygmas are globular (Fig. 6K; Fig. 13E, H). Pollen of A. pungens and A. ledebourii is tricolporate, oblong-spheroidal, elliptical in equatorial view, roundedtrilobed in polar view, with distinct, deep, and long colpi (ectoapertures). Atraphaxis pungens from Askiz, Khakassia (255) has syncolporate pollen 29.3–32.4 × 20.5–24.5 µm, A. ledebourii (sub A. pungens) from Kharalyg-Khem, Tuva (247) has the same pollen size (27.7 (25.8–31.8) × 26.3 (24.5–27.9) µm (Yurtseva et al. 2014). The pollen surface of A. pungens has striato-perforate ornamentation with distinct, protruding, intensively branching and merging striae and grooves with 2–7 pits or perforations in a row (Fig. 7C). Atraphaxis pungens 304 from Kartushibinskii ridge has spherical pollen grains 30 µm in diameter with abnormal flat ridges and slit-like grooves (not shown). Thus, A. pungens and A. ledebourii do not show reliable morphological characteristics which let to clearly distinguish them. Due to high morphological similarity of A. pungens and A. ledebourii, their putative hybrid A. pungens × A. ledebourii (243) from Kaa-Khem village (Tuva, Kyzylskii Raion) does not differ from the parents. This hybrid has larger leaves at vegetative shoots, larger perianths and fruits (Fig. 10F) The reciprocal hybrid A. ledebourii × A. pungens 318 from Hangay, Mongolia, has black, shiny fruits common for A. pungens. Three specimens of A × uyukensis (241, 301, 311) from Kara-Khaak and the Uyuk ridge, Tuva, combine plastid haplotype U and ITS ribotypes P2 and P2del of A. pungens (303, 316, 317, 319) (Figs 1, 3). The sample (301) from Kara-Khaak, Tuva (Fig. 9F) resembles other specimens of A. pungens in the appearance, long vegetative shoots, broadly elliptical leaf blades, perianth, and black glossy fruits (Fig. 14A–C). The samples 241 and 311 (Fig. 11) resemble the samples A. pungens (184, 192, 304) from Kartushibinskii ridge (Fig. 9A–C) in the appearance, oblongoblanceolate leaf blades of short generative and vegetative shoots, small thyrses, and yellowish perianths (Fig. 14D–F). Their stygmas are flat, which is a characteristic of A. pungens (compare Fig. 12D–F and Fig. 14G–I)., Published as part of Yurtseva, Olga V., Vasilieva, Natalia V., Kostikova, Vera A. & Samigullin, Tahir H., 2022, A broadly sampled 3 - loci plastid phylogeny of Atraphaxis (Polygoneae, Polygonoideae, Polygonaceae) reveals new taxa: III. A. kuvaevii and сryptic species in A. pungens from Southern Siberia and Northern Mongolia, pp. 13-63 in Phytotaxa 566 (1) on pages 25-33, DOI: 10.11646/phytotaxa.566.1.2, http://zenodo.org/record/7103418, {"references":["Yurtseva, O. V., Severova, E. E. & Bovina, I. Yu. (2014) Pollen morphology and taxonomy of Atraphaxis (Polygoneae, Polygonaceae). Plant Systematics and Evolution 300 (4): 749 - 766. http: // doi. org / 10.1007 / s 00606 - 013 - 0917 - 4"]}

Atraphaxis kuvaevii sp. nov. Atraphaxis kuvaevii, a local endemic of Tuva, forms a separate subclade in both plastid and ITS trees and does not show introgression with other species (Figs 1 and 3). It is a dwarf schrub 10–25 cm in height, with divaricately branched second-year shoots and annual generative and vegetative shoots 2–5 cm long (Fig. 4, Table 1). Leaves are small, rhomboid (Fig. 5A). Generative shoots terminate with thyrses with 3–5 cymes of 1–2 flowers. The perianth is small, with a tube articulated to a pedicel in ochreola (Fig. 5B, C). The epidermal cells covering the perianth tube and the tepals do not differ from those of A. pungens and A. ledebourii (Fig. 6A–F). Fruits are smooth and glossy, light brown, with three styles terminated with globular stigmas (Fig. 5D–F, Fig. 6G–I). Atraphaxis kuvaevii has tricolporate pollen grains 27.4–27.7 × 14.5–15.9 µm with striate-perforate sporoderm ornamentation: it has short grooves and 2–3 perforations in a row (Fig. 7A–B), while the pollen surface of A. pungens (Fig. 7C) and A. decipiens (Fig. 7D–E) has distinct striae and long grooves with 5–6 perforations in a row. Atraphaxis kuvaevii clearly differs from other species of Atraphaxis with the smallest leaf blades, perianth, and the tepals cuneate at the base, the filiform perianth tube articulated to the pedicel in ochreolae, and the smallest fruits (Table 1). Atraphaxis kuvaevii was previously mistaken for A. laetevirens distributed in Kazakhstan, Altai and Dzhungaria. Both species have small glands at the edges and on the lower surface of leaf blades, and light-brown glossy fruits, but the fruits of A. kuvaevii are equal to the inner tepals, while the fruits of A. laetevirens are 2–3 times smaller than the inner tepals of the perianth. Atraphaxis laetevirens falls different subclades in both plastid and ITS trees. Thus, the morphological and molecular data support A. kuvaevii as a distinct species distributed in Tuva and the South of Krasnoyarskii Krai (Fig. 8). RUSSIA. Krasnoyarskii Krai: the West Sayan, Sayano-Shushensky Nature Reserve, left board of the Sayano-Shushenskii water reservoir, above the Kalbakmys river, stony mountain steppe, 615 m, 3 July 1987, Kuvaev & Sonnikova 1015-13 (holotype MW0061320), Published as part of Yurtseva, Olga V., Vasilieva, Natalia V., Kostikova, Vera A. & Samigullin, Tahir H., 2022, A broadly sampled 3 - loci plastid phylogeny of Atraphaxis (Polygoneae, Polygonoideae, Polygonaceae) reveals new taxa: III. A. kuvaevii and сryptic species in A. pungens from Southern Siberia and Northern Mongolia, pp. 13-63 in Phytotaxa 566 (1) on pages 21-22, DOI: 10.11646/phytotaxa.566.1.2, http://zenodo.org/record/7103418

2. Atraphaxis ledebourii Yurtseva, sp. nov. Type: — RUSSIA. Tuva: Tes-Khem Raion, the southern slope of the East Tannu-Ola ridge, dry stone bed of the Khyralyg-Khem river, 1200 m, 3 June 1972, Khanminchun 4821 (holotype MW0061283!, isotypes LE!, MHA!, NSK!) (Fig. 21). Description: —Shrub ca 30–50 cm tall. Woody shoots up to 1 cm in diam., inclined-spreading, dark-gray, much branched, bark exfoliating and fibrously disintegrating, naking pinkish or creamy wood. Second-year shoots bear lateral branchlets 5–12 cm, 3–5 mm in diam. departing at 45 degrees, somewhat spiny after fruiting, but easily break off. Annual shoots 3–15 cm long, 0.5–1.0 mm in diam., straight, finely ribbed, glabrous, creamy, leafy, with internodes 5.0– 6.5 mm. Some annual shoots terminate with frondo-bracteose thyrses 1–2 cm long with 3–8 congested (rarely far spaced) cymes of 2–3 flowers. Leaf blades bright-green, later bluish-green or gray, rhomboid-elliptical, broadly elliptical, or obovate, 7–12(30) × 5–7(18) mm, obtuse, shortly pointed, or acuminate, suddenly narrowed to a petiole 1.0–1.0 mm, glabrous, with prominent net of veins, with smooth, flat or slightly undulate margin. Leaf ratio 2–3. Ochreas 4–7 mm long, semitransparent, silvery-whitish, cleft in two linear-lanceolate lacinulas 3–5 mm long, each with yellowish or white vein, and middle truncated lacinula finely inciso-serrated at top. Lateral lacinulas are fibrously disintegrated at margins. Ochreolas 2.5–4.0 mm, membranous, light-brown at base, transparent above, cleft in two sharp semitransparent subulate lacinulas at both sides of strongly reduced leaf blade. Perianth white, creamy, or yellowish, with filiform tube 2.0– 5.5 mm joined to a pedicel 1.5–4.0 mm long with articulation. Outer tepals rounded or elliptical, 2.0–4.0 × 2.0–4.0 mm, prominently reticulate, reflected to a pedicel. Inner tepals rotundate, 4.0–5.5(7) × 4.0–5.0 (7) mm, with slightly crenulate margin and reticulate venation. Stamens 8, filaments subulate-filiform, suddenly dilated at base. Anthers rounded-oval. Achenes 2.8–4.5 × 1.4–2.0 mm, 1.5 times shorter than inner tepals of perianth, ovoid, suddenly acuminate, trigonous, with almost flat faces, obtuse ribs, smooth, glossy, black. Styles three, 0.4–0.5(0.7) mm long, fused to 1/4–2/5, with fimbriate-capitate stigmas 0.3–0.4 mm in diam. Etymology: —named after Ledebour who reported this taxon from Altai as Tragopyrum pungens M.Bieb. Flowering time: —June. Fruiting time: —July–August. Distribution: MONGOLIA. [Uvs aimag, Zavkhan aimag]: Ubsunur depression, North Hangay; RUSSIA. Tuva: the East Tannu-Ola ridge, the Uyuk ridge along the Yenisey River; Altai Republic. Ecology: —rocks, dry gravely and steppe slopes. Chromosome numbers: —from 42 to 72, RUSSIA. Altai Republic: Kosh-Agach Raion, vicinity of Kokorya vill., Kostikov & Kostikova (NSK3000910!), and RUSSIA. Republic of Tuva: Chaa-Khol’ Raion, the shore of the SayanoShushenskii reservoir, Kostikov & Kostikova (NSK3000907!), both sub A. pungens (M. Bieb.) Ledeb. (Voronkova et al. 2019)., Published as part of Yurtseva, Olga V., Vasilieva, Natalia V., Kostikova, Vera A. & Samigullin, Tahir H., 2022, A broadly sampled 3 - loci plastid phylogeny of Atraphaxis (Polygoneae, Polygonoideae, Polygonaceae) reveals new taxa: III. A. kuvaevii and сryptic species in A. pungens from Southern Siberia and Northern Mongolia, pp. 13-63 in Phytotaxa 566 (1) on pages 46-48, DOI: 10.11646/phytotaxa.566.1.2, http://zenodo.org/record/7103418, {"references":["Voronkova, M. S., Banaev, E. V., Tomoshevich, M. A. & Kostikova, V. A. (2019) IAPT chromosome data 30 / 12. In: Marhold, K. et al. (ed.) IAPT chromosome data 30. Taxon 68 (5): 1130, E 29."]}

1. Atraphaxis kuvaevii O.V.Yurtseva, sp. nov. Type: — RUSSIA. Krasnoyarskii Krai: West Sayan Mountains, Sayano-Shushenski Nature Reserve, left board of the Sayano-Shushenskaya water reservoir, above the Kalbakmys River, stony mountain steppe, 615 m, 3 July 1987, Kuvaev & Sonnikova 1015-13 (holotype MW0061320!, isotype NS!) (Figs 4–5, 6G–I, 7A–B). Description (macromorphology): —Dwarf shrub 15–20(30) cm tall with tortuous extensively branched shoots. Second-year shoots 3–4 mm in diam, smooth, slightly prickly, covered with dark gray exfoliating bark hiding foxybrown wood. Annual vegetative shoots to 10–12 (15) cm long, straight, creamy, finely ribbed, glabrous, 1.0 mm in diam., leafy, with internodes 5–6 mm long. Generative shoot 1.5–5 cm long, with 3–4 leaves spaced by internodes 1–2 mm long, terminates with bracteose thyrse 10–15 mm long, with 3–4 cymes of 1–2(3) flowers. Ochreas at vegetative shoots 3–6 mm long, membranous, slightly brownish at base, above semitransparent, silvery-white at margin, split in 2 subulate lacinulas serrate-incised at margin, each with a single faintly visible vein and short middle lacinula serrateincised at margin. Ochreolas in thyrses 2,5– 3 mm long, semitransparent, cleft in two sharp teeth. Leaf blades dark green, bright green or glaucose, 8–11 × 2–6 mm at shoots, 3–4 × 1 mm in thyrses, elliptical or rhomboid-elliptical, apex rounded or shortly pointed, base cuneate, gradually or suddenly narrowed into a petiole 0.5–1.0 mm, thick, leathery, with a clear net of veins, glabrous above, finely crenulate and papillate at margin (Fig. 5A). Perianth ca. 5–6 mm long; filiform part of perianth tube 2.0–3.0 mm long, with a funnel-form extention 0.3–0.4(0.5) mm long; the tube is much longer than a pedicel 1.0 mm, or the place of articulation is hidden in ochreola (Fig. 5B–D). Outer tepals two, 2.0–2.5 × 1.5–2.0 mm, rhomboid, ovate or rotundate, reflected to a pedicel; inner tepals three, 3.0–4.0 × 2.5–2.0 mm, flat, base cuneate, apex rotundate, green at the middle, with a wide pinkish margin, slightly undulate, surround the achene, with prominent longitudinal veins or without veins. Stamens 8, filaments subulate-lanceolate, gradually dilated towards the base, anthers rounded-oval. Achene 2.0–2.8 × 1.3–1.6 mm, equal to inner tepals, trigonous, ovoid, gradually acuminate, light-brown, smooth, glossy, faces ovate- or elliptical-lanceolate, almost flat, ribes obtuse, styles three 0.3–0.4 mm long, free from the base, with papillate globular stigmas 0.2 mm (Figs 5E–F, 6G–H). Description (pollen): —Oblong-spheroidal, 24.2–27.3 × 19.3–20.7 µm (P/E = 1.29); tricolporate, elliptical in equatorial view, rounded-trilobed in polar view; colpi are distinct, long and deep; ora distinct, lalongate or circular (Fig. 7A, B). The sporoderm ornamentation is striate-perforate with the striae 0.10–0.20 µm thick, distinct and protruding, mainly short and straight, divided by deep short grooves with 2–4 (rarely more) large perforations in a row. Exine is 2.0 (1.7–2.4) µm thick. Etymology: —Named after Vladimir B. Kuvaev (1918—2009), a researcher of Siberian Flora who first indicated diagnostic characteristics of the taxon. Flowering time:— June–July. Fruiting time:— July–August. Distribution:— RUSSIA. Southern Siberia, Krasnoyarskii Krai: West Sayan Mts, valley of the Yenisey river in Tuvinian Depression, rocky slopes of the Uyuk ridge and the West and East Tannu-Ola ridge facing the Yenisey river, Khayyrakan Mt. Ecology:— Stony mountain steppe, nanophyton steppe, feather grass & selaginella shrubby steppe, granite or limestone rocks and slopes. Note on the holotype: —MW0061320 was previously identified as: A. pungens (M.Bieb.) Jaub. & Spach (12 March 1988, det. Kuvaev.), A. laetevirens (Ledeb.) Jaub. & Spach (23 March 1988, det. Kuvaev), A. frutescens (L.) K.Koch (23 Mart 1988, det. Lomonosova); A. frutescens (L.) K.Koch (20 April 1988, det. Kuvaev), A. frutescens (L.) K.Koch × A. laetevirens (Ledeb.) Jaub. & Spach (6 March 2015, det. Yurtseva). Specimens examined:— RUSSIA. Krasnoyarskii Krai: West Sayan Mts, Sayano-Shushensky Nature Reserve, left board of Sayano-Shushenskaya water reservoir, above the mouth of the creek Taldy-Chas, base of granit slope, 540 m, 3 July 1987, Kuvaev & Sonnikova 1013-6 (MW0061321!); Republic of Tuva [Ulug-Khemskii Raion]: the eastern slope of Mt Khayyrakan, on the main pass of the main saddle, on rocks. 6 July 1946, Schroeter 820 (MW0061323!); ibidem (MHA!); ibidem, 11 June 1981, Doronkin (NSK); ibidem, vicinity of Khayyrakan village, the steep bank of the Yenisey River, N51°34’27.0’’, E93°03’58.8’’, 11 July 2014, Kostikov (NSK!); ibidem, Mt Khayyrakan, stony steppe, 13 July 1974, Timokhina & Egorova 528 (NSK!); West Sayan Mts, the Uyuk ridge, environs of Bayan-Kol vill., southern slope, eastern microslope, feather grass & selaginella shrubby steppe, 7 July 1975, Lomonosova & Grushevskya 2708 (MHA!); ibidem, southern slope, eastern microslope, at rocks, 8 July 1975, Lomonosova & Iskakova 2709 (MHA!); ibidem (NSK!); 10 km E of Shagonar vill., the Shagonar rocks on the left bank of the Yenisey River, limestone rocks on the banks of the Yenisey, 650 m, 51°34’N, 93°04’E, 17 August 2002, Nikitin, Byalt & Sytin 1120 (LE!); valley of the Yenisey river, vicinity of Iiji-Tal settlment, 51°34’05.9”N, 93° 23’16.4”E, 19 July 2006, Shmakov et al. K3-2360 (ALTB!); [Biy-Khem kozhuun at the border with the Krasnoyarskii Krai], the right board of the Sayano-Shushenskaya hydropower reservoir, the Sayan canyon at the mouth of the Chamge (Chinge) river, flat surface of the ancient upper terrace of the Yenisey river, nanophyton steppe with Atraphaxis and Spirea hypericifolium dominated, at the upper sides of deep ravines, 540–600 m, N51°42’39.9’’, E092°17’46.5’’, 17 July 2014, Sonnikova (MW!); [Kyzyl kozhuun] right bank of the Yenisey river 40 km West of Kyzyl, at the rock “Dzharga”, in a ravine on the southern gravelly slope, 27 July 1947, Schroeter 281 (MHA!)., Published as part of Yurtseva, Olga V., Vasilieva, Natalia V., Kostikova, Vera A. & Samigullin, Tahir H., 2022, A broadly sampled 3 - loci plastid phylogeny of Atraphaxis (Polygoneae, Polygonoideae, Polygonaceae) reveals new taxa: III. A. kuvaevii and сryptic species in A. pungens from Southern Siberia and Northern Mongolia, pp. 13-63 in Phytotaxa 566 (1) on pages 45-46, DOI: 10.11646/phytotaxa.566.1.2, http://zenodo.org/record/7103418

Key to the species of Atraphaxis growing in Mongolia, Altai, Southern Siberia and Russian Transbaikalia 1. Perianth with two inner and two outer tepals, 6 stamens, lenticular fruit with two styles................................................................ 2 — Perianth with three inner and two outer tepals, 8 stamens, trigonous fruit with 3 styles................................................................. 4 2. Leaf blades shortly puberulent abaxially. Altai Mts, Zaisan........................................................................................ A. canescens — Leaf blades glabrous......................................................................................................................................................................... 3 3. Inner perianth tepals reniform, 7–9 × 9–11 mm, 1.5–2 times longer than fruits. Altai Mts, Zaisan.............................. A. compacta — Inner perianth tepals rotundate, 5–6 × 7–8 mm, almost equal to fruits........................................................................... A. replicata 4. Outer perianth tepals 1.5 times shorter than inner tepals, outspread in fruiting. All perianth tepals depart from a cup-shaped extention of a filiform perianth tube 0.7–1.0 mm long joint to a pedicel 2.0–3.0 mm long...................................... A. selengensis — Outer perianth tepals 2–3 times shorter than inner tepals, reflected to a pedicel in fruiting. All the tepals depart from the top of a wedge-shaped extention of a filiform perianth tube 2.5–6.0 mm long, equal to a pedicel.............................................................. 5 5. Subshrub or shrub with orthotropic generative shoots terminated with thyrse 5–25 cm long, sometimes branching, never prickly after fruiting...................................................................................................................................................................................... 6 — Shrub with sturdy and prickly second-year shoots and axillary branchlets bearing lateral generative shoots with compact thyrses 1.5–4 cm long................................................................................................................................................................................... 8 6. Shrub up to 100 cm tall. Generative shoot terminates with branched thyrse 10–25 cm long with well spaced cymes of flowers. Leaf blades broadly elliptical..................................................................................................................................................... A. virgata — Subshrub 20–70 mm tall. Generative shoot terminates with thyrse 5–12 cm long with 8–12 congested cymes of flowers, sometimes with paracladia. Leaf blades oblanceolate or linear-lanceolate........................................................................................................ 7 7. Leaf blades oblanceolate, with net of veins, more than 3 mm in width. Ochreolas transparent...... A. frutescens subsp. frutescens — Leaf blades linear-lanceolate, with a midvein beneath, less than 3 mm in width, ochreolas herbaceous.......................................................................................................................................................................................................... A. frutescens subsp. sibirica 8. Annual shoots and edges of leaf blades shortly papillose. Fruits light-brown................................................................................. 9 — Annual shoots and leaf blades glabrous. Fruits black or dark-brown............................................................................................ 10 9. Leaf blades bright-green, rotundate to broadly elliptical, 7–15 × 8–10 mm. Tepals white or creamy, inner tepals 5–6 × 6–7 mm. Perianth tube is twice longer than a pedicel. Achene 2–3 times shorter than inner tepals. Kazakhstan, Altai Mts, Saur, Tarbagatay, Dzungaria..................................................................................................................................................................... A. laetevirens — Leaf blades dark-green, rhomboid-ovate to rhomboid-elliptical, 8–11 × 2–6 mm. Tepals dark-green with pink margin, inner tepals 3–4 × 2–4 mm. Perianth tube articulated to a pedicel in ochreola.Achene is equal or slightly shorter than inner tepals. Krasnoyarskii Krai, Tuva......................................................................................................................................................................... A. kuvaevii 10. Ochreas 7–9(12) mm long. Fruits black, lanceolate, finely pitted or microtuberculate (× 4), glossy, equal to inner tepals of perianth. Styles fused to 1/3–2/5, 0.6–0.8 mm long, stygmas spongy, campaniform in flowering. Transbaikalia........................ A. davurica — Ochreas 3–7 mm long. Fruits black or dark-brown, ovoid to lanceolate, smooth, glossy, twice shorter or equal to inner tepals of perianth. Styles fused to 1/4–1/5, 0.3–0.5 mm long, stygmas globular or flat, in flowering bent to ovary. Altai Mts, Khakassia, Krasnoyarskii Krai, Tuva, Mongolia.............................................................................................................................................. 11 11. Leaf blades broadly elliptical to oblanceolate, gradually narrowed to a petiole, leaf ratio 3–7. Inner tepals of perianth 4–6 × 4–6 mm. Fruits black and glossy, ovoid, 2.4–3.3 × 1.4–2.0 mm, twice shorter than inner tepals 4–6 × 4.5–6 mm. Krasnoyarskii Krai, Khakassia, Tuva, Mongolia.............................................................................................................................................. A. pungens Note: Hybrids of A. × uyukensis (the Uyuk ridge) have black ovoid fruits with microtuberculate surface, 4.1 × 2.0 mm. Hybrids A. pungens × A. frutescens subsp. sibirica have dark-brown lanceolate fruits 2.8 × 1.3 mm with distinct ribs. — Leaf blades rhomboid-elliptical, broadly elliptical, or obovate, leaf ratio 2–3. Fruits dark-brown to black, 2.8–4.5 × 1.4–2.0 mm, 1.5 times shorter than inner tepals of perianth, which are 5–7 × 5–7 mm. Mongolia, Altai Mts, Tuva....................... A. ledebourii Note: Hybrids Atraphaxis ledebourii × A. frutescens subsp. sibirica have dark-brown lanceolate fruits 4.5 × 2.0 mm with distinct ribs., Published as part of Yurtseva, Olga V., Vasilieva, Natalia V., Kostikova, Vera A. & Samigullin, Tahir H., 2022, A broadly sampled 3 - loci plastid phylogeny of Atraphaxis (Polygoneae, Polygonoideae, Polygonaceae) reveals new taxa: III. A. kuvaevii and сryptic species in A. pungens from Southern Siberia and Northern Mongolia, pp. 13-63 in Phytotaxa 566 (1) on pages 44-45, DOI: 10.11646/phytotaxa.566.1.2, http://zenodo.org/record/7103418

Elwendia lindbergii There are relatively a few species of Elwendia distributed in the Tajik Gorny Badakhshan. In the latest floristic treatment for Tajikistan (Korovin et al. 1984), only three species were reported from this territory. Two of them, Elwendia persica and E. badachschanica (Kamelin 1977: 190) Pimenov & Kljuykov in Degtjareva et al. (2013: 1002), are common in this region and well recognizable. The third species, E. chaerophylloides, more rarely distributed in this region, is also easily distinguished from the aforementioned species due to its remarkable fruits, separating into mericarps from the upper part. Elwendia lindbergii was described from Afghan Badakhshan (Rechinger & Riedl 1963) and insufficiently studied. On the boundary territory to Tajik Gorny Badakhshan, there are specimens morphologically very similar to E. lindbergii. These plants are very similar in their habit and possess identical leaf blades with refracted rachis, umbelles, bracts and bracteoles. The fruits of the specimens from Tajik Gorny Badakhshan separate into mericarps from the base like in the majority of species of Umbelliferae including Elwendia lindbergii. We propose to treat the critical specimens of Elwendia from Tajik Gorny Badakhshan as Elwendia lindbergii. According to our molecular phylogenetic analyses, two specimens of E. lindbergii from this region do not group together and fall into unresolved position with E. intermedia, E. chaerophylloides and E. persica. Although both specimens of E. lindbergii contain slightly different ITS/ETS sequences, they are morphologically quite homogeneous. We believe the nucleotide sequence variability of E. lindbergii can be caused by the geographical isolation of their populations under harsh climatic conditions of Gorny Badakhshan. With this record, the distribution area of Elwendia lindbergii significantly expands to the east; and the species is reported as new to Tajikistan and Central Asia (Fig. 4). ITS and ETS data do not allow to differentiate E. lindbergii from E. intermedia and E. chaerophylloides due to unresolved relationships between different accessions of these species. In contrast, morphological characters allow easy identification of E. lindbergii, distinguished from E. intermedia by solid (vs. hollow) stems, longer terminal segments of radical leaves (3–5 mm vs. 2–3 mm), a lower number of umbel rays (2–7 vs. 10–23) and a lower number of bracteoles (2–5 vs. 6–10). These characters suggest that E. lindbergii should be retained as a distinct species., Published as part of Kljuykov, Eugene V., Lazkov, Georgii A., Degtjareva, Galina V., Zakharova, Ekaterina A., Ukrainskaja, Uljana A., Lyskov, Dmitry F. & Samigullin, Tahir H., 2022, Taxonomic revision of critical taxa of Elwendia (Umbelliferae-Apioideae) from Tajikistan and neighbouring countries, with descriptions of three new species from the Pamir-Alay and distributional novelties, pp. 255-273 in Phytotaxa 543 (5) on page 268, DOI: 10.11646/phytotaxa.543.5.1, http://zenodo.org/record/6479422, {"references":["Korovin, E. P., Pimenov, M. G. & Kinzikayeva, G. K. (1984) Umbelliferae Moris. In: Ovchinnikov, P. N. (Ed.) Flora of the Tajik SSR, vol. 7. Academy of Sciences of USSR, Moscow & Leningrad, pp. 10 - 214. [In Russian]","Kamelin, R. V. (1977) New taxa of Central Asian Umbelliferae. Novosti Sistematiki Vysshikh Rastenij 14: 187 - 192.","Degtjareva, G. V., Kljuykov, E. V., Samigullin, T. H., Valiejo-Roman, C. M. & Pimenov, M. G. (2013) ITS phylogeny of Middle Asian geophilic Umbelliferae-Apioideae genera with comments on their morphology and utility of psbA-trnH sequences. Plant Systematics and Evolution 299: 985 - 1010. https: // doi. org / 10.1007 / s 00606 - 013 - 0779 - 9","Rechinger, K. H. & Riedl, H. (1963) Umbelliferae. In: Koeie, M. & Rechinger, K. H. Symbolae afghanicae 5. Biologiske Skrifter udgivet af det Kongelige Danske Videnskabernes Selskab 13 (4): 27 - 135."]}

Elwendia capusii is new to Afghanistan Elwendia capusii is a species widely distributed in Mountain Middle Asia, Tian-Shan and Pamir-Alay. In South Tajikistan, it occurs up to the Afghan border but has not been reported from Afghanistan (Rechinger 1987). Based on herbarium specimens examined at E, K, MW, M and LE, we confirm the presence of the species in North Afghanistan. Elwendia capusii is morphologically distinctive among the other species due to the presence of long (over a half of the mericarp length) and erect (vs. recurved) styles. Our phylogenetic tree highlights that the E. capusii clade is well delineated and supported, but also a genetic differentiation among its populations has occurred. In the absence of any good distinguishing morphological characters, the observed variation in ITS/ETS sequences between the samples from Afghanistan and from the remaining area can be considered as infraspecific. Specimens examined:— Afghanistan, Mashad to Faizabad Badakhshan (Kochka valley), 3500 ft., limestone slopes. 19 May 1964, F.Furse 6147 (K!, LE!). Afghanistan, prov. Balkh: Ali Koh, 5 km SO Sultan Baba (18 km S Mazar-i-Sharif), 1200 m, Kalkfelsen. 67/09-36/34. 11 April 1978, Podlech 31015 (M!); prov. Balkh, Mazar-i-Scharif, 29 May 1969, Hedge, Wendelbo & Ekberg 8485A (E!, MW!). Distribution area:— Kazakhstan, Kyrgyzstan, Uzbekistan, Turkmenistan, Tajikistan, N Afghanistan., Published as part of Kljuykov, Eugene V., Lazkov, Georgii A., Degtjareva, Galina V., Zakharova, Ekaterina A., Ukrainskaja, Uljana A., Lyskov, Dmitry F. & Samigullin, Tahir H., 2022, Taxonomic revision of critical taxa of Elwendia (Umbelliferae-Apioideae) from Tajikistan and neighbouring countries, with descriptions of three new species from the Pamir-Alay and distributional novelties, pp. 255-273 in Phytotaxa 543 (5) on page 268, DOI: 10.11646/phytotaxa.543.5.1, http://zenodo.org/record/6479422, {"references":["Rechinger, K. H. (1987) Bunium. In: Hedge, I. C., Lamond, J. M., Rechinger, K. H., et al. (1987) Umbelliferae. In: Rechinger, K. H. (Ed.) Flora iranica, vol. 162. Akademische Druck- und Verlagsanstalt, Graz, Austria, pp. 236 - 257."]}

The Elwendia kuhitangi – E. salsa complex in the Pamir-Alay Elwendia kuhitangi was described by Nevski (1937) from the Kuhitang Range as Bunium kuhitangi Nevski (1937: 274). In the protologue, a detailed description of mature fruits that is required for certain identification is lacking. In floristic publications, all high-mountain dwarf plants occurring in the Pamir-Alay from the Kuhitang and Western Hissar Ranges to Darwaz (Korovin 1950, 1959; Kljuykov & Pimenov 1983; Korovin et al. 1984) and in North Afghanistan and East Iran (Rechinger 1987) passed under the name B. kuhitangi. Elwendia salsa was first described as Bunium salsum Korovin (1924: 86) from the vicinities of Samarkand and treated as closely related to Elwendia kuhitangi by Kljuykov (1988). In contrast to E. kuhitangi, E. salsa is morphologically more fully characterised and has a broader distribution area in midlands and low mountains of the Western Pamir-Alay and Western Tian-Shan. Kljuykov (1988) formally described Bunium subsect. Salsa to accommodate E. kuhitangi and E. salsa. The shared characters used to justify this taxonomy were as follows: fruits with a strong smell, hardly separating into mericarp, pedicels length equal to a fruit; any of these characters is not unique in Elwendia. According to our results, E. kuhitangi and E. salsa represent a distinct lineage, placed in a clade together with E. ugamica. Thus, the morphology-based classification was confirmed by our phylogenetic analysis. Based on the present study, we expand the description of E. kuhitangi with fruit characters and propose diagnostic characters to distinguish between E. kuhitangi s.str. and E. salsa s.str., such as the length of terminal segments of the uppermost stem leaves (3–5 mm vs. 7–12 mm), number of umbel rays (2–3 vs. 5–10), shape and size of mericarps (narrowly linear vs. linear). Besides, the fruits of E. kuhitangi separate easier into mericarps and have a less expressed smell. With regards to the distribution, E. kuhitangi is restricted to the Kuhitang Range (Tukmenistan, Uzbekistan) (Kljuykov et al. 2018), whereas E. salsa is characterised by a broader distribution range from Uzbekistan and Tajikistan to Kyrgyzstan (Korovin 1959; Kljuykov & Pimenov 1983; Korovin et al. 1984; Kljuykov et al. 2018)., Published as part of Kljuykov, Eugene V., Lazkov, Georgii A., Degtjareva, Galina V., Zakharova, Ekaterina A., Ukrainskaja, Uljana A., Lyskov, Dmitry F. & Samigullin, Tahir H., 2022, Taxonomic revision of critical taxa of Elwendia (Umbelliferae-Apioideae) from Tajikistan and neighbouring countries, with descriptions of three new species from the Pamir-Alay and distributional novelties, pp. 255-273 in Phytotaxa 543 (5) on pages 267-268, DOI: 10.11646/phytotaxa.543.5.1, http://zenodo.org/record/6479422, {"references":["Nevski, S. A. (1937) Materials for flora of Kuhitang and its foothills. Transactions of Institute of Botany of Akademy of Science of SSSR. Series 1, flora and systematics of higher plants 4: 199 - 346. [In Russian]","Korovin, E. P. (1950) Bunium L. In: Schischkin, B. K. (Ed.) Flora of USSR, vol. 16. Academy of Sciences of the URSS, Moscow & Leningrad, pp. 396 - 411. [In Russian]","Korovin, E. P. (1959) Umbelliferae. In: Vvedensky, A. I. (Ed.) Flora of Uzbekistan, vol. 4. Academy of Science Uzbek SSR, Tashkent, pp. 257 - 470. [In Russian]","Kljuykov, E. V. & Pimenov, M. G. (1983) Bunium L. In: Adylov, T. A. (Ed.) Opredelitel' rasteniy Sredney Asii: kriticheskiy konspekt flory, vol. 7. Fan, Tashkent, pp. 228 - 235. [In Russian]","Korovin, E. P., Pimenov, M. G. & Kinzikayeva, G. K. (1984) Umbelliferae Moris. In: Ovchinnikov, P. N. (Ed.) Flora of the Tajik SSR, vol. 7. Academy of Sciences of USSR, Moscow & Leningrad, pp. 10 - 214. [In Russian]","Rechinger, K. H. (1987) Bunium. In: Hedge, I. C., Lamond, J. M., Rechinger, K. H., et al. (1987) Umbelliferae. In: Rechinger, K. H. (Ed.) Flora iranica, vol. 162. Akademische Druck- und Verlagsanstalt, Graz, Austria, pp. 236 - 257.","Korovin, E. P. (1924) Umbelliferae turkestanicae novae. Botanicheskie materialy Gerbariya Glavnogo Botanicheskogo Sada RSFSR 5 (6): 81 - 86.","Kljuykov, E. V. (1988) A survey of the genus Bunium L. Revision of the generic system. Byulleten' Moskovskogo Obshchestva Ispytatelei Prirody. Otdel biologicheskii 93: 76 - 89. [In Russian]","Kljuykov, E. V., Tojibaev, K. Sh. & Lyskov, D. F. (2018) The genus Elwendia Boiss. (Apiaceae) in the flora of Uzbekistan. Turczaninovia 21 (2): 210 - 220. https: // doi. org / 10.14258 / turczaninowia. 21.2.18"]}

The Chinese endemic Polygonum schischkinii was studied from both morphological and molecular points of view. On the basis of the structure of ocreas and ocreolas, P. schischkinii appears to be a member of the genus Atraphaxis. The results of Maximum Likelihood and Bayesian analyses of combined data of the plastid [rpl32-trnL(UAG) IGS, trnL(UAA) intron, trnL-trnF IGS] and nuclear rDNA ITS1-2 regions, carried out on 61 members of tribe Polygoneae, confirmed position of P. schischkinii in the Atraphaxis clade. A new combination-Atraphaxis glareosa-based on P. glareosum, which has nomenclatural priority over P. schischkinii, is proposed. Ecological notes and a detailed distributional map of the species are also provided. © 2021 Magnolia Press. All rights reserved.