Key to the varieties of Artemisia sacrorum
Artemisia sacrorum Ledeb., Mem. Acad. Imp. Sci. 5:571 (1805)
A. messerschmidtiana Besser, Nouv. Mém. Soc. Imp. Naturalistes Moscou 3: 27 (1834).
A. sacrorum var. minor f. discolor Kom., Fl. Manschur. 3(2): 664 (1907).
A. gmelini var. discolor (Kom.) Nakai, Fl. Kor. 2: 31 (1911).
A. freyniana f. discolor (Kom.) Kitag., J. Jap. Bot. 41: 367 (1936).
A. sacrorum subsp. manshurica Kitam., Acta Phytotax. Geobot. 7: 66 (1938).
A. sacrorum var. messerschmidtiana (Besser) Y. R. Ling, Bull. Bot. Res., Harbin 8(4): 13 (1988).
Subshrubs. Roots thin fibrous. Rhizomes long creeping, woody. Stems 52–108 cm tall, 1.9–3.5 mm thick; indumentum silky pubescent while young, sparsely arachnoid-pubescent later. Lower leaves withering before flowering. Middle leaves petiolate 14.9–21.4 mm long; leaf blades triangular-ovate, elliptic-ovate, bipinnately divided, 43.1–65.1 mm long, 39.1–52.7 mm wide, sparsely villous pubescent with white glandular on adaxial surface, densely villous pubescent on abaxial surface; segments 5–7 pairs, lanceolate, pinnately divided, 19.1–25.3 mm long, 7.1–11.6 mm wide; ultimate segments linear-lanceolate, triangular pectinate; rachis triangular pectinate. Upper leaves small, leafy bracts, simple. Inflorescence broad panicle. Capitula long pedunculate, nodding; involucre subglobose, 2.3–2.7 mm long, 1.9–2.4 mm wide; phyllaries 4–5 seriate, laxly imbricate, densely villous; outer phyllaries ovate-lanceolate, green, narrowly membranous margin, acute at apex; middle phyllaries lanceolate, green, middle membranous margin, obtuse at apex; inner phyllaries obovate, broadly membranous margin, rounded at apex. Receptacle conical, glabrous with glandular. Marginal female florets 9–14 in number, corolla 0.8–1.1 mm long, conical with cuneate shaped glandular, 2–3 toothed irregularly at apex; style 0.8–1.3 mm long, longer than corolla; stigma 0.5–0.7 mm long, 2-divergent, somewhat emarginate at apex; ovary fertile, 0.5–0.8 mm long. Disk bisexual florets 7–11 in number, corolla 1.3–1.6 mm long, narrowly tubular without throat, cuneate shaped glandular, 5-toothed regularly at apex; style 0.8–1.3 mm long, longer than corolla; stigma 0.4–0.6 mm long, 2-divergent, combed funnelform at apex; ovary fertile, 0.5–0.8 mm long; anthers 5 in number, 2 acute basal appendages, apical appendage acute, triangular, 0.5–0.7 mm long.
Type: Russia. Tibet, Sibir (type: ?).
Flowering: Sep. to Oct.
Distribution: China, Russia, Korea, Afghanistan, Mongolia.
Distribution of Korea: Hamgyeongbuk-do, Yanggang-do, Hwanghae-do.
Habitat: Hills, waysides.
Note:
A. sacrorum Ledeb var.
sacrorum can be distinguished from other related taxa by small subshrub, non-glandular trichome eliminated at flowering time on adaxial leaf surface, white glandular trichome distributed on adaxial leaf surface, non-glandular trichome of white-gray on abaxial leaf surface and phyllaries (
Fig. 1). It has the same taxon as
A. messerschmidtiana treated as
A. sacrorum var.
messerschmidtiana by
Ling (1988). However,
A. messerschmidtiana needs to be treated as a synonym of
A. sacrorum because its priority and characters with densely villous on only abaxial leaf surface. According to specimens, this taxon is restrictively distributed from Siberia to Manchuria including North Korea. It is not distributed in South Korea or Japan (
Fig. 5).
Kitamura (1937) has published
A. sacarorum subsp.
manshurica with the size of capitula, the shape of a leaf, and geographical delimitation by comparing to
A. sacrorum of Altai region and to the illustration (Icones Pl. fl. Ross. tab. 471) of
Ledebour (1834) without considering the geographical variation through observation of limited specimens. In Korea, the scientific name of
A. freyniana f.
discolor combined by Kitagawa (1996) is used. However, this scientific name was published by
Komarov (1907) without considering geographical variations. Therefore, we treated those names as synonyms of
A. sacororum including
A. gmelini var.
discolor by
Nakai (1911).
Artemisia sacrorum Ledeb.
var. incana (Besser) Y. R. Ling, Bull. Bot. Res., Harbin 8(4): 13 (1988). (
Fig. 2)
A. messerschmidtiana var. incana Besser, Nouv. Mém. Soc. Imp. Naturalistes Moscou 3: 27 (1834).
A. sacrorum f. vestita Kom., Fl. Manschur. 3(2): 664 (1907).
A. gmelini var. vestita (Kom.) Nakai, Fl. Kor. 2: 31 (1911).
A. freyniana f. vestita (Kom.) Kitag., J. Jap. Bot. 41: 367 (1936).
A. sacarorum var. vestita Kitam., Acta Phytotax. Geobot. 7: 66 (1938).
Subshrubs. Roots thin fibrous. Rhizomes long creeping woody. Stems 56–105 cm tall, 1.5–3.9 mm thick; indumentum silky pubescent while young, sparsely arachnoid-pubescent later. Lower leaves withering before flowering. Middle leaves petiolate 13.1–19.1 mm long; leaf blades triangular-ovate, elliptic-ovate, bipinnately divided, 41.6–61.4 mm long, 41.1–59.1 mm wide, densely villous pubescent with glandular on both surfaces; segments 5–7 pairs, lanceolate, pinnately divided, 21.9–28.5 mm long, 8.2–11.2 mm wide; ultimate segments linear-lanceolate, triangular pectinate; rachis triangular pectinate. Upper leaves small, leafy bracts, simple. Inflorescence broad panicle. Capitula long pedunculate, nodding; involucre subglobose, 2.7–3.0 mm long, 1.9–2.7 mm wide; phyllaries 4–5 seriate, laxly imbricate, densely villous with glandular; outer phyllaries ovate-lanceolate, narrowly membranous margin, acute at apex; middle phyllaries lanceolate, middle membranous margin, obtuse at apex; inner phyllaries obovate, broadly membranous margin, rounded at apex. Receptacle conical, glabrous with glandular. Marginal female florets 5–10 in number, corolla 0.6–1.2 mm long, conical with cuneate shaped glandular, 2–3 toothed irregularly at apex; style 0.7–1.3 mm long, longer than corolla; stigma 0.5–0.8 mm long, 2-divergent, somewhat emarginate at apex; ovary fertile, 0.5–0.9 mm long. Disk bisexual florets 8–13 in number, corolla 1.2–1.7 mm long, narrowly tubular without throat, cuneate shaped glandular, 5-toothed regularly at apex; style 0.6–1.2 mm long, longer than corolla; stigma 0.3–0.5 mm long, 2-divergent, combed funnel form at apex; ovary fertile, 0.5–0.8 mm long; anthers 5 in number, 2 acute basal appendages, apical appendage acute, triangular, 0.4–0.6 mm long.
Type: Sibir (type: ?).
Flowering: Sep. to Oct.
Distribution: China, Korea, Mongolia.
Distribution of Korea: Hamgyeongbuk-do, Yanggang-do.
Habitat: Hills, waysides, shrublands, slopes, often dominant on S slopes, roadsides, forest steppes.
Note: This taxon was recognized as a new species of
A. messerschmidtiana by
Besser (1834) at first with two infraspecific taxa: α.
viridis, β.
incana. He treated
A. sacrorum as a synonym of α.
viridis (=
A. sacrorum var.
sacrorum) characterized by glabrous adaxial leaf surface and white-gray trichome on abaxial leaf surface. Also, β.
incana was first described with characters of white-gray trichome on both leaf surfaces. After that,
Ling (1988) classified α.
viridis and β.
incana as varieties of
A. sacrorum because of their morphological similarities. We also found out that there was no difference among them except the distribution pattern of trichome between
A. sacrorum and α.
viridis, β.
incana. This taxon is considered to have geographical distribution from Siberia to Manchuria, including North Korea. However, it is not distributed in South Korea or Japan (
Fig. 5). In Korea, the scientific name of
A. freyniana f.
vestita combined by Kitagawa (1996) is used. However, the name of
A. sacrorum var.
minor f.
vestita published by
Komarov (1907) is a combination without considering geographical variations. We treated
A. freyniana f.
vestita as a synonym of
A. sacororum var.
incana including
A. gmelini var.
vestita suggested by
Nakai (1911) and
A. sacarorum subsp.
manshurica var.
vestit used by
Kitamura (1937).
Artemisia sacrorum Ledeb.
var. iwayomogi (Kitam.) M. S. Park & G. Y. Chung, comb. et stat. nov. (
Figs. 3–
4).
Basionym: A. iwayomogi Kitam., Acta Phytotax. Geobot. 7: 65 (1938).
A. sacrorum subsp. iwayomogi (Kitam.) Vorosch., Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 102(1): 68 (1997).
A. messerschmidtiana f. laxiflora Nakai, 2: 31 (1911).
A. sacrorum subsp. laxiflora (Nakai) Kitag., Neo-Lineam. Fl. Mansh. 431 (1979).
Subshrubs. Roots thin fibrous. Rhizomes long creeping woody. Stems 71–153 cm tall, 2.8–4.2 mm thick; indumentum silky pubescent while young, sparsely arachnoid-pubescent later. Lower leaves withering before flowering. Middle leaves petiolate 13.3–20.3 mm long; leaf blades triangular-ovate, elliptic-ovate, bipinnately divided, 53.7–105.4 mm long, 40.8–67.2 mm wide, sparsely villous with glandular on both surfaces; segments 5–8 pairs, lanceolate, pinnately divided, 24.5–35.8 mm long, 8.3–12.4 mm wide; ultimate segments linear-lanceolate, triangular pectinate; rachis triangular pectinate. Upper leaves small, leafy bracts, simple. Inflorescence broad panicle. Capitula long pedunculate, nodding; involucre subglobose, 2.7–3.1 mm long, 2.7–3.0 mm wide; phyllaries 4–5 seriate, laxly imbricate, sparsely villous with glandular; outer phyllaries ovate-lanceolate, narrowly membranous margin, acute at apex; middle phyllaries lanceolate, middle membranous margin, obtuse at apex; inner phyllaries obovate, broadly membranous margin, rounded at apex. Receptacle convex, glabrous, rarely a few T-shaped trichome with glandular. Marginal female florets 5–12 in number, corolla 0.7–1.1 mm long, conical with cuneate shaped glandular, 2–3 toothed irregularly at apex; style 0.7–1.0 mm long, longer than corolla; stigma 0.4–0.6 mm long, 2-divergent, somewhat emarginate at apex; ovary fertile, 0.5–0.8 mm long. Disk bisexual florets 5–8 in number, corolla 1.3–1.5 mm long, narrowly tubular without throat, cuneate shaped glandular, 5- toothed regularly at apex; style 0.9–1.3 mm long, longer than corolla; stigma 0.3–0.5 mm long, 2-divergent, combed funnelform at apex; ovary fertile, 0.5–0.7 mm long; anthers 5 in number, 2 acute basal appendages, apical appendage acute, triangular, 0.5–0.7 mm long. Achenes brown, pyriform, 1.2–1.5 mm long, glabrous. Chromosome number 2n = 54.
Type: Japan. Hokkaido, Prov. Shiribeshi, Setana, 13 Oct. 1935. S. Kitamura (type: in KYO!).
Flowering: Sep. to Oct.
Distribution: China, Russia, Japan, Korea, Afghanistan, N India, Kazakhstan, Mongolia, Nepal, N Pakistan.
Distribution of Korea: All provinces.
Habitat: Hills, waysides, shrublands, slopes, often dominant on slopes, roadsides, forest steppes.
Note:
A. gmelinii has been used as a correct name for this taxon (
Lee, 1996). After reviewing the original description and illustration of
Besser (1834) and Ledebour (1805;
1833) through typification and the illustration (Gmel. Fl. Sib. No. 106, Tab. LVI.
Fig. 1),
A. gmelinii is widely distributed in Siberia. However, it is not distributed in the Far East (
Maximowicz; 1872,
Komarov; 1907;
Boyko, 1990).
Kitamura (1937) has published
A. iwayomogi (nomen novum) instead of a wrong scientific name that some scholars have misused. It was distinguished from
A. sacrorum of Altia region. He mentioned that
A. sacrorum had small sized plant and leaf, acute leaflet, and definitely different geographical distribution. He described that
A. iwayomogi was distributed in Japan, Sakhalin, Kuril Islands, the Korean peninsula, and the Far East of Manchuria. After observing specimens of Russia, China, and Japan,
A. gmelinii is mainly distributed in Siberia but not the Far East, while
A. sacrorum var.
sacrorum (=
A. messerschmidtiana) and
A. sacrorum var.
incana have expanded their distribution from Ural mountains to Dahuria of the northeast, Far East, the northeast China, and North Korea. In addition,
A. sacrorum var.
iwayomogi is distributed in Mongolia, northeast China, Japan, Sakhalin, Kuril Islands, the Far East of Manchuria, and the entire Korean peninsula including the area mentioned by
Kitamura (1937). According to the original description and type specimen of
A. iwayomogi published by
Kitamura (1937), the typification of
A. sacrorum var.
iwayomogi distributed in South Korea is the same. The distribution of
A. sacrorum var.
iwayomogi almost overlaps with
A. sacrorum var.
sacrorum and
A. sacrorum var.
incana. Considering that there is no difference except that
A. iwayomogi has no non-glandular trichome on both leaf surface or phyllaries of capitula at flowering time,
A. iwayomogi needs to be treated as a variety of
A. sacrorum.
The chromosome number of
A. sacrorum var.
sacrorum and
A. sacrorum var.
incana has been reported to be 2
n = 2
x = 18 (
Kaul and Bakshi, 1984;
Volkova and Boyko, 1985). The reported chromosome number of
A. sacrorum var.
iwayomogi is 2
n = 6
x = 54 (
Park et al., 2009). Inferring evolution trend from geographical distribution, morphological characters, and chromosome numbers of the
A. sacororum group, when
A. sacrorum var.
sacrorum and
A. sacrorum var.
incana (diploid) with similar geographical distribution moved to the Far East, the Korean Peninsula, and Japan, the trichome might have been eliminated from both leaf surfaces to adapt to the environment. At that time,
A. sacrorum var.
iwayomogi (haxaploid) might have been divided (
Fig. 5). Such hypothesis about their relationship has been supported by
Park and Chung (2012). They have reported that the pattern of pollen surface,
A. sacrorum var.
sacrorum, and
A. sacrorum var.
incana has a connected base of spinule on pollen. However,
A. sacrorum var.
iwayomogi has a disconnected base of spinule on pollen.
Consequently, there are three varieties under A. sacrorum in the Korean peninsula: A. sacrorum var. sacrorum with white-gray non-glandular trichome on adaxial leaf surface, A. sacrorum var. incana with white-gray non-glandular trichome on both leaf surfaces, and A. sacrorum var. iwayomogi with almost glabrous on both leaf surfaces. A. sacrorum var. sacrorum and A. sacrorum var. incana are distributed mainly in Siberia, expanding from Ural mountains to Dahuria of the northeast, Far East of Russia, northeast China, and North Korea, while A. sacrorum var. iwayomogi is widely distributed in Mongolia, northeast China, Japan, Sakhalin, Kuril Islands, the Far East of Manchuria, and the Korean Peninsula.