Atriplex pseudotatarica Sukhor. & Nidhan Singh, 2023

Sukhorukov, Alexander P., Singh, Nidhan, Kushunina, Maria, Zaika, Maxim A. & Sennikov, Alexander N., 2023, A new species of Atriplex (Amaranthaceae) from the Indian subcontinent, PhytoKeys 229, pp. 167-183 : 167

Atriplex pseudotatarica Sukhor. & Nidhan Singh sp. nov.

Fig. 1

Atriplex crassifolia - Atriplex crassifolia auct. non C.A.Mey.

Atriplex leucoclada - Atriplex leucoclada auct. non Boiss.

Atriplex - Atriplex spp. div. in herb. DD and K.

Type.

India. Haryana, surroundings of Panipat town, near Asan Kalan village , 29°15.1286'N, 76°31.4816'E, 15 Nov 2022 [in flowering and early fruiting stages], N. Singh & A. Sukhorukov 9 (Holotype: CAL, isotype: BSD) .

Description.

Monoecious subshrub up to 1.5 m high, branched in upper half; leaves alternate, shortly petiolate; petioles up to 1.0 cm long; blades greyish-silvery on both sides, 1.0-4.0 × 0.5-1.0 cm (much smaller towards inflorescence), oblong or narrowly oblong, entire or shallowly sinuate, with Kranz-anatomy; inflorescences branched, up to 15 cm long, with pseudopposite bracts or with a few small leaves forming pseudowhorls (after fruiting turning into small alternate leaves in younger shoots), aphyllous in other parts; glomerules condensed or slightly interrupted, of both male and female flowers, the latter are also located at the axils of uppermost leaves below the main inflorescence; male flowers stipitate at base, with 5 free perianth segments, anthers 0.25 mm long; bract-like cover of female flowers (Fig. 2A, B ) rhombic, entire or scarcely dentate, with or without lateral angles, smooth at the back or rarely with 1-2 very short outgrowths, valves connate to the halfway, sometimes to one third of their length, with indistinct veins, heteromorphic in some other characters: (1) bract-like cover of female flowers located in leaf axils ± indurated in lower half and inflated at fruiting, 4.5-5.5 mm long, rhombic, and (2) bract-like cover of female flowers located in the main inflorescence slightly indurated and not inflated at fruiting, 2.0-4.0 mm long, trilobate and rhombic; seeds heterospermic (Fig. 2C, D ): seeds developing in fruits located below the main inflorescence black, slightly elongated (1.1-1.3 × 1.4-1.6 mm), ripening earlier (in November; obs. in Haryana State, India) compared with those of the fruits located in the main inflorescence; seeds in fruits located in the main inflorescence black (similar to those developing below the main inflorescence) or yellowish-brown, 0.8-1.0 mm in diameter, ripening in late November-December.

Phenology.

Flowering: July-November; fruiting: November-December.

Habitat.

Saline soils, sands, wasteland, roadsides, 0-2200 m a.s.l. In the natural landscapes in Haryana, Atriplex pseudotatarica was observed together with Bassia indica (Wight) A.J.Scott, Suaeda fruticosa Forssk. (all - Amaranthaceae ), and some grasses.

Etymology.

The specific epithet is chosen due to the resemblance of the new species to A. tatarica L., which also has long aphyllous inflorescences.

Conservation status.

Although there is currently a limited number of collected specimens of Atriplex pseudotatarica , this species is clearly more overlooked than rare. Given that it is often found in disturbed habitats, produces a large number of seeds and is naturally adapted to saline substrates, we propose that the species should be assigned to the IUCN Red List category "Least Concern" ( IUCN Standards and Petitions Committee 2022).

Distribution

(Fig. 3 ). India, Pakistan and SE Afghanistan.

Specimens examined.

Afghanistan. [Laghman province] Alingar valley , 6000 ft, 1 Sep 1956, W. Thesiger 1693 (BM) .

India. Delhi, 23 Oct 1874, anonymous 23395 (K); [Himachal Pradesh] Kimawar [Kinnaur], 1884, J.R. Drummond 535 (DD-29978); [Haryana], Karnal, 1885, J.R. Drummond 26479 (K) ; Haryana, surroundings of Panipat town, nr Asan Khurd village , 29°18.1584'N, 76°31.8779'E, 15 Nov 2022, N. Singh & A. Sukhorukov 7 (CAL, BSD) .

Pakistan. Lahore, 1846, T.T. Thomson s.n. (K); [Punjab] Rawalpindi, 1872, J.E.T. Aitchinson 224 (K); [Gilgit-Baltistan prov.] Skardu , 7000-8000 ft, 15 Jul [18]92 [early flowering], without collector’s name 12060 (DD!); [Khyber Pakhtunkhwa prov., Swat Distr.] Shohdara , 11 Nov 1935 [in fruiting stage], R.R. Stewart 15362 (DD-77925); Lahore, common in weedy places, 17 Oct 1938 [in flowering stage], Parker s3436 (DD-81928, DD-81929, K); [Punjab province], nr Attock, 15 Nov 1956, R.R. Stewart 27830 (K) .

Notes.

All examined herbarium specimens of A. pseudotatarica are represented by upper twigs of the plants. They were mostly incorrectly identified as A. crassifolia , or rarely left without identification, as Atriplex sp. To date, the name A. crassifolia may be found misapplied to some other species attributable to different groups of the genus. Atriplex crassifolia is an annual C3-species belonging to A. sect. Teutliopsis Dumort. ( Moser 1934; Iljin 1936; Sukhorukov 2006; Žerdoner Čalasan et al. 2022) with a restricted distribution range in semideserts of Kazakhstan and South-West Siberia, Russia ( Iljin 1936; Sukhorukov 2006), penetrating into western China (Sukhorukov in Nobis et al. 2016). All other records of A. crassifolia reported from Europe are erroneous ( Sukhorukov 2006; Sukhorukov et al. 2019). Aellen (1939), Ivanov (1989) and Medvedeva (1996) erroneously applied this name to A. patens (Litv.) Iljin, another species from A. sect. Teutliopsis ( Sukhorukov 2006). The specimens from the Mediterranean area (GBIF Sekretariat 2022; re-identifications in BM!, K!, LE!) belong to A. tornabenei Tineo (C4-clade, A. sect. Obionopsis (Lange) Dumort.: Sukhorukov et al. 2022). The name A. crassifolia has also been widely used for the plants growing in lowlands of India and Pakistan (e.g., Hooker (1890), Bamber (1916)), and it is still erroneously applied in recent floras, checklists and ecological studies ( Puri et al. 1964; Shetty and Singh 1991; Hussain and Mirza 1993; Jain et al. 2000; Kumar 2001; Paul 2012; Kumar and Singh 2013; Ibrahim 2019). Hooker (1890) stated that A. crassifolia is present in both lowlands ("westwards of Jumna [Yamuna] River") and high mountains. Nevertheless, all records of A. crassifolia from the Himalayas refer to C4-species from A. sect. Obione (Gaertn.) Reichenb.: A. pamirica Iljin and A. schugnanica Iljin [= A. pallida (Moq.) Sukhor. ≡ Chenopodium pallidum Moq., nom. rejic. prop.], and those from the lowlands and foothills belong to A. pseudotatarica .

Some of the plants growing in the lowlands of Pakistan also belong to A. pseudotatarica , of which some specimens were misidentified as A. leucoclada Boiss. Hedge (1997) noted that this species is highly polymorphic in the area covered in "Flora Iranica", with the extreme forms having smooth, apically trilobate bract-like cover. Unfortunately, he did not indicate where such specimens were collected, but such characters are typical of A. pseudotatarica .

Phylogenetic relationships

(Fig. 4 ). Based on the combined nrITS and nrETS analyses, A. pseudotatarica was found sister to A. schugnanica , and both form a subclade within the Eurasian clade, A. sect. Obionopsis (Lange) Dumort., which comprises ~ 15 C4-species predominantly distributed in Irano-Turanian floristic region ( Sukhorukov et al. 2022; Žerdoner Čalasan et al. 2022). Atriplex pseudotatarica and A. schugnanica share some characters typical of the members of A. sect. Obionopsis (aphyllous inflorescences, sclerified bract-like cover with the valves connate up to the half of their length, presence of the female flowers in leaf axils and both female and male flowers in the inflorescence), but have several conspicuous morphological differences (Table 3 ). Additionally, A. pseudotatarica is distributed in the lowlands and pre-Himalayan foothills, whereas A. schugnanica is a typical montane plant growing in the West Himalayas, Karakoram and Pamir at the altitudes of (2000) 2600-4800 m a.s.l. ( Iljin 1936; Sukhorukov et al. 2019). In Table 3 , we also included other similarly looking C4 Atriplex species; three of them (except A. tatarica ) are present in the lowlands of Pakistan, and only one ( A. pseudotatarica ) is reaching India. Atriplex tatarica , widely distributed in many parts of the "Flora Iranica" area, as well as A. kalafganica Aellen (Aellen in Podlech 1975) are also added here because of their morphological resemblance with A. pseudotatarica .

The most conspicuous characters of A. pseudotatarica are subshrubby life form, pseudopposite leaves or leaves in pseudowhorls below the inflorescence seen at fruiting, and presence of black seeds.

Figures

Figure 1. Atriplex pseudotatarica. A an overview of the plant B a twig with the inflorescence C a vegetative shoot D a shoot at fruiting. Origin of the material A Haryana, near Asan Khurd village, Nov 2022 B Haryana, near Asan Kalan village, Nov 2022 C Haryana, near Asan Kalan village, Aug 2022 D Haryana, near Panipat town, Oct 2014. Photographer: A, B A. Sukhorukov, C, D N. Singh.

Figure 2. Bract-like cover (A, B) and seeds (C, D) of A. pseudotatarica A cover of a female flower located below the main inflorescence B cover of a female flower located in the main inflorescence C black seed D yellowish-brown seed. Scale bars: 1 mm.

Figure 3. Records of A. pseudotatarica.

Figure 4. Maximum clade credibility tree from the BEAST analysis of the ITS + ETS Atriplex dataset. Bayesian posterior probabilities are given above the branches, bootstrap percentages of the maximum likelihood analyses are given below the branches.