Delphinium lacostei Danguy (1908: 50)

Li, Hui-Min, Yuan, Qiong & Yang, Qin-Er, 2018, Taxonomic studies on the genus Delphinium (Ranunculaceae) from China (XVI): Three new synonyms of D. lacostei, Phytotaxa 382 (2), pp. 151-166 : 163-166

Delphinium lacostei Danguy (1908: 50) . Figs. 1–15 .

Type:— PAKISTAN. Sasser-La, 4950 m, 7 September 1906, Lacoste s.n. (holotype P!). Fig. 15A .

= Delphinium tetragynum W.T. Wang (1983: 26) , syn. nov.

Type:— CHINA. Xinjiang: Taxkorgan, Mintaka Pass , 4590 m, 6 July 1978, Northwest Inst. Bot. Xinjiang Exped. 1119 (holotype PE!; isotypes WUK!, XJBI!). Fig. 1 .

= Delphinium yechengense C.Y. Yang & B. Wang (1992: 82) , syn. nov.

Type:— CHINA. Xinjiang: Yecheng, Kokyar Pass, gravelly slopes, 3800 m, 31 July 1984, Abulimit & Abulikim 137 (holotype XJA!). Fig. 3A .

= Delphinium taxkorganense W.T. Wang (1993: 349) , syn. nov.

Type:— CHINA. Xinjiang: Taxkorgan, Khunjerab Pass, gravelly slopes near glacier, 4700 m, 12 August 1989, S.K Wu, Y.H. Wu & Y. Fei 4891 (holotype KUN!). Fig. 4A .

Description:—Perennial. Stems often in clumps, erect, usually branched, 9–35 (–60) cm tall, sparsely to densely spreading whitish puberulent or subglabrous. Basal leaves 3–9 (–12), long petiolate; petiole (2–) 6–14 (–21) cm long, subglabrous or sparsely puberulent, narrowly sheathed; blade glabrous or subglabrous adaxially, sparsely pubescent on nerves abaxially, (1.5–) 3–7.5 cm long, 2–6 (–12) cm wide, orbicular-cordate or reniform-cordate, base cordate, 3-parted nearly to the middle, median partition cuneate or broadly obovate, indistinctly 3-lobate, lobes mucronate, lateral partitions obliquely flabellate, unequally 2-lobate. Middle stem leaves absent or 1, occasionally 2, smaller, shortly petiolate; petiole 1.5–4.5 (–8) cm long; blade 1.5–4.8 cm long, 2–4 (–6) cm wide, deeply 3-divided almost to the base, lobules ovate. Distal stem leaves bractlike. Inflorescence racemose-corymbose, (1–) 2–6-flowered, (3–) 7–20 (–30) cm long; peduncle densely spreading whitish puberulent; bracts leaflike, trilobate or entire, 1–4.5 cm long, sparsely puberulent. Pedicels 4–8 (–14) cm long, densely spreading whitish puberulent; bracteoles far or less distant from flowers, sometimes subtending them, linear or oblong-linear, 7–11 (–18) mm long, 1.5–3 mm wide, sparsely puberulent. Sepals blue or blue-purple, abaxially whitish puberulent, persistent; upper sepal 9–18 mm long, 6–10 mm wide, obtuse, the spur saccate-conic, obtuse at apex, slightly recurved, 6–12 (–20) mm long, 7–11 mm wide at base; other sepals 11–20 mm long, 9–18 mm wide. Petals dark brown or black, acuminate, glabrous or sparsely ciliate. Staminode dark brown, limb oblong, 9–12 mm long, 3–5 mm wide, 2-cleft and gaping almost halfway, the lobes yellow barbate and white ciliate, claw 4.5–6 mm long. Filaments sparsely pilose or glabrous. Carpels usually 4–5, very rarely 6; ovaries sparsely to densely puberulent. Follicles oblong, ca. 1.5 cm long, densely puberulent. Seeds ca. 1.2 mm long, dark brown, cylindrical, transversely squamulose.

Distribution and habitat:— Delphinium lacostei is distributed in eastern Afghanistan, western China (Xinjiang), northern India, and northern Pakistan ( Fig. 16 ). It often grows on gravelly or grassy slopes or among boulders at altitudes of 3500–4800 m above sea level. It was recorded to occur in Kyrgyzstan ( Wang & Warnock 2001), but we have not as yet seen any material from that country.

Phenology:—Flowering July to August; fruiting September.

Additional specimens examined:— AFGHANISTAN. Badakshan: Wakhan, C. Grey-Wilson & T.F. Hower 1449 (E), C. Grey-Wilson & T.F. Hower 1541 (E, K), S. Penn 13 (K); Kunar: Nari, D. Podlech 16432 (K), D. Podlech 16504 (K).

CHINA. Xinjiang: Pamir (precise locality unknown), Alexeenko 428 (LE), Alexeenko 429 (LE), Haiying s.n. (XJA); Taxkorgan, Y.S. Chen 13–2276 (PE), N.R. Cui 22 (XJA), N.R. Cui 23 (XJA), N.R. Cui 24 (XJA), N.R. Cui 25 (XJA), N.R. Cui 169 (XJA), J.J. Feng 70043 (XJA), H.M. Li & Q.G. Mao 339 (IBSC), H.M. Li & Q.G. Mao 341 (IBSC), H.M. Li & Y.P. Zeng 655 (IBSC), Northwest Inst. Bot. Xinjiang Exped. 1195 (PE, WUK), Northwest Inst. Bot. Xinjiang Exped. 1264 (WUK, XJBI), Northwest Inst. Bot. Xinjiang Exped. 1470 (WUK, XJBI), Northwest Inst. Bot. Xinjiang Exped. 1495 (PE, WUK, XJBI), Pan-Himalaya Flora Exped. FPH-1380 (PE), J. Qiu & J.J. Feng 106 (XJA), J. Qiu & J.J. Feng LiuJQ0106 (KUN), H.Y. Ren 35 (XJA), S.K. Wu, Y.H. Wu & Y. Fei 5037 (KUN, PE); Yecheng, R.F. Huang 124 (HNWP), B.S. Li, Z.Q. Du & B.N. Zhang 11428 (PE), B.S. Li, Z.Q. Du & B.N. Zhang 11479 (PE), Qinghai- Xizang Exped. 1465 (KUN, PE).

INDIA. Himachal Pradesh: Chamba, J.H. Lace 1655 (E), J.H. Lace 2094 (E); Kinnaur, G. Sherriff 7460 (E); Lahaul and Spiti, R. McBeath 2151 (E), R. McBeath 2153 (E), G. Watt. 2495 (E).

PAKISTAN. Khyber Pakhtunkhwa: Kohistan, O. Polunin 6208 (E), O. Polunin 6250 (BM, E).

Notes:— Delphinium lacostei and D. brunonianum are somewhat confused with each other because of close similarity in their spur shape. The former is distinguishable from the latter mainly by the smaller flowers with the upper sepal 9–18 mm long, 6–10 mm wide (vs. 20–30 mm long, almost as wide) and the stem, peduncle and pedicel all whitish non-glandular (vs. densely yellowish glandular) pubescent. Moreover, we have not noticed any musky odor in D. lacostei , a feature of D. brunonianum mentioned in the literature and often written on the collection labels. Geographically D. lacostei seems to be restricted in the Pamir region while D. brunonianum is essentially a Himalayan species. In China, D. brunonianum is currently known only from Xizang (Tibet).

The morphology of the primarily Kashmir Himalayan Delphinium cashmerianum Royle (1834: 55) and its relationship with D. lacostei is worthy of a further study. From the herbarium material of D. cashmerianum we have seen, this species seems to be quite closely similar to D. lacostei in general appearance, although Munz (1967) keyed out D. lacostei from D. cashmerianum by the sepal length (16 mm vs. 20–30 mm long) and described D. cashmerianum as having strigose or spreading pubescent pedicels and 3–7 follicles. Field observations of D. cashmerianum is badly needed.

Figures

FIGURE 1. Holotype (A) and isotype (B−D) sheets of Delphinium tetragynum (= D. lacostei). Inset in Figure 1B: puberulent carpels.

FIGURE 2. Paratype (A) and isoparatype (B−D) sheets of Delphinium tetragynum (= D. lacostei).

FIGURE 3. Specimens of Delphinium lacostei. A. CHINA. Xinjiang:Yecheng, Kokyar Pass, Abulimit & Abulikim 137 (XJA, holotype of D. yechengense). B−D. CHINA. Xinjiang:Yecheng, B.S. Li, Z.Q. Du & B.N. Zhang 11479 (PE). Inset in Figure 3B: 4 carpels in the flower arrowed; inset in Figure 3D: 5 carpels in the flower arrowed.

FIGURE 4. Holotype sheet (A) of Delphinium taxkorganense (= D. lacostei) and the line illustration (B) of D. taxkorganense given in the protologue.

FIGURE 5. Specimens of Delphinium lacostei. A. CHINA. Xinjiang: Taxkorgan, Dangbashi, Northwest Inst. Bot. Xinjiang Exped. 1495 (PE). B, C. Same locality, Northwest Inst. Bot. Xinjiang Exped. 1495 (WUK). D. Same locality, Northwest Inst. Bot. Xinjiang Exped. 1495 (XJBI). Note that the PE sheet was referred to D. lacostei by Wang (1983), but annotated as D. yechengense by him in 1995 on the determination slip.

FIGURE 6. Delphinium lacostei in the wild (CHINA. Xinjiang: Taxkorgan, Mintaka Pass, the type locality of D. tetragynum). A. Habitat. B. Habit. C. Rhizome. D. Stems. E. Leaves (adaxial surface). F. Inflorescences. G. Peduncle and pedicels. H. Pedicel.

FIGURE 7. Delphinium lacostei in the wild (CHINA. Xinjiang: Taxkorgan, Mintaka Pass, the type locality of D. tetragynum). A. Flowers and pedicels with bracts and bracteoles arrowed. B. Staminodes. C. Petals. D. Stamens. E. Carpels. F. Follicles. G. Seed.

FIGURE 8. Ten individuals within a population of Delphinium lacostei (CHINA. Xinjiang: Taxkorgan, Mintaka Pass, the type locality of D. tetragynum), showing the variation in plant size and flower number.

FIGURE 9. Trichome morphology of pedicels (A−D) and carpels (E−H) in a population of Delphinium lacostei (CHINA. Xinjiang: Taxkorgan, Mintaka Pass, the type locality of D. tetragynum). A, E. Mixture of basally swollen, rough-surfaced and filiform, smoothsurfaced trichomes (× 35). B, F. Mixture of basally swollen, rough-surfaced and filiform, smooth-surfaced trichomes (× 300). C, G. Basally swollen, smooth-surfaced trichomes (× 2000). D, H. Filiform, rough-surfaced trichome (× 2000).

FIGURE 10. Carpels in 12 individuals within a population of Delphinium lacostei (CHINA. Xinjiang: Taxkorgan, Mintaka Pass, the type locality of D. tetragynum), showing the variation in number and pubescence. The ratio of carpel number/flower number is given for each individual.

FIGURE 11. Delphinium lacostei in the wild (CHINA. Xinjiang: Taxkorgan, Khunjerab Pass, the type locality of D. taxkorganense). A. Habitat. B. Habit. C. Flowers and pedicels with bracts and bracteoles arrowed. D. Staminodes. E. Petals. F. Stamens. G. Carpels. H. Follicles. I. Seed.

FIGURE 12. Eight individuals within a population of Delphinium lacostei (CHINA. Xinjiang: Taxkorgan, Khunjerab Pass, the type locality of D. taxkorganense), showing the variation in plant size and flower number.

FIGURE 13. Trichome morphology on pedicel (A−D) and carpel (E−H) of Delphinium lacostei (CHINA. Xinjiang:Taxkorgan, Khunjerab Pass, the type locality of D. taxkorganense). A, E. Mixture of basally swollen, rough-surfaced and filiform, smooth-surfaced trichomes (× 35). B, F. Mixture of basally swollen, rough-surfaced and filiform, smooth-surfaced trichomes (× 300). C, G. Basally swollen, smoothsurfaced trichomes (× 2000). D, H. Filiform, rough-surfaced trichome (× 2000).

FIGURE 14. Carpels in nine individuals within a population of Delphinium lacostei (CHINA. Xinjiang: Taxkorgan, Khunjerab Pass, the type locality of D. taxkorganense), showing the variation in number and pubescence. The ratio of carpel number/flower number is given for each individual.

FIGURE 15. Specimens of Delphinium lacostei. A. PAKISTAN. Sasser La, Lacoste s.n. (holotype, P). B. AFGHANISTAN. Badakshan: Wakhan, C. Grey-Wilson & T.F. Hower 1541 (E; misidentified as D. brunonianum on the determination slip). C. INDIA. Himachal Pradesh: Chamba, Pangi, J.H. Lace 2094 (E). D. PAKISTAN. Khyber Pakhtunkhwa: Kohistan, O. Polunin 6208 (E).

FIGURE 16. Distribution of Delphinium lacostei (●).