Dactylostalix ringens Reichenbach (1878: 74)

Suetsugu, Kenji, Yagame, Takahiro, Yukawa, Tomohisa & Suyama, Yoshihisa, 2024, Taxonomic revision of the formerly monotypic orchid genus Dactylostalix, Phytotaxa 652 (2), pp. 83-99 : 89-95

Dactylostalix ringens Reichenbach (1878: 74) ( Figs. 1A–C , 2 , 7B–C & 9–11 ).

Type:— JAPAN. Kinki District— Kii (= Wakayama Pref.: Mt. Koya?), June 1875, Rein s.n. (holotype: W0196072!).

= Dactylostalix maculosa Miyabe & Kudo (1915: 3) . Type:— JAPAN. Hokkaido District —Oshima Prov.: Kamiiso-gun, Moheji-mura (= Hokkaido Pref.: Hokuto-shi, Moheji), 16 May 1906, S. Ninoue s.n. (lectotype: SAPS036659!), designated here.

= Dactylostalix ringens Reichenbach (1878: 74) var. punctata Miyabe & Tatewaki (1937: 49) ≡ Dactylostalix ringens Reichenbach (1878: 74) f. punctata ( Miyabe & Tatewaki 1937: 49) Yonekura (2011: 239) . Type:— JAPAN. Hokkaido District — Kushiro Prov.: Kamioboro (= Hokkaido Pref.: Akkeshi-gun, Akkeshi-cho, Kamioboro), 14 June 1937, M. Tatewaki s.n. (holotype: SAPS036661!).

= Calypso japonica Maximovich in Komarov (1901: 533) ≡ Calypso bulbosa ( Linnaeus 1753: 951) Oakes (1842: 28) var. japonica (Maximovich in Komarov 1901: 533) Makino (1905: 14). Type:— JAPAN. Chubu District— Fujiyama (= Shizuoka Pref.: Fujinomiya-shi, Mt. Fuji), 1894, Tschonoski (= Chonosuke Sukawa) s.n. (lectotype: the third individual from the right in the upper row of LE 01012254!), designated here.

Rhizome creeping, pale brown. Scape arising from the rhizome, erect, (10–) 15–26 cm long, 1.5–2.5 mm in diameter at the middle, glabrous, pale green, bearing one flower at the apex. Leaf solitary; leaf blade elliptic to ovate, 37–78 mm long, 20–46 mm wide, pale green, occasionally with reddish-purple spots, obtuse at the apex, rounded or obtuse at the base; petiole broadly winged, 10–28 mm long. Bract 2.1–3.2 mm long, pale green. Ovary and pedicel 6.5–10.4 mm long, pale green. Dorsal sepal oblanceolate to oblong, 18–30 × 3.3–7.5 mm, yellowish-green with reddish-purple spots around the base, apex obtuse or acute. Lateral sepals obliquely oblong to falcate, 16–28 × 2.6–5.7 mm, yellowish-green with reddish-purple spots around the base, apex obtuse or acute. Lateral petals oblanceolate to oblong, 16–27 × 3.2–7.2 mm, yellowish-green with reddish-purple spots around the base, apex obtuse or acute. Labellum protruding, distinctly reflexed at the apex, 3-lobed, 12–19 × 9.5–13.9 mm, disc with 2 low keels from base to basal 1/3; lateral lobes slightly obliquely semiorbicular, 5.0–9.3 × 2.8–4.0 mm, white with a large reddish-purple spot at the apex of each lobe; posterior portion of midlobe ovate to cordate, 7.8–13.2 × 9.5–13.9 mm, white with randomly arranged small reddish-purple spots and 2 larger reddish-purple dots at the base, margin slightly undulate and irregularly erose-denticulate. Column curved, 9.7–13.2 × 3.4–7.0 mm at the widest part (a little above the middle) and 1.9–3.5 mm wide at the base, semi-cylindrical with column wings at the lateral part of the apex and 1 appendage at the central part of the apex; rostellum well-developed; stigma located just below the rostellum, 2.0–4.5 × 1.8–4.2 mm. Anther hemispheric, 1.3–2.0 mm in diameter; pollinia 4, in 2 pairs, obovoid, 1.3–2.0 × 0.8–1.1 mm; viscidium rhomboid, 0.6–1.0 mm long. Capsule obovoid, 2.2–2.9 cm long.

Additional specimens examined:— RUSSIA. Kuril Islands— Etorofu Island: Mt. Ooyama , 28 July 1931, Ohwi s.n. ( SAPS) . JAPAN. Hokkaido District — Hokkaido Pref.: Akkeshi-gun, Akkeshi-cho, Kamioboro , 14 June 1937, Tatewaki 28156 ( SAPS) ; loc. cit., 14 June 1937, Tatewaki 28156 ( SAPS) ; loc. cit., 14 June 1937, Tatewaki 28156 ( SAPS) ; loc. cit., 14 June 1937, Tatewaki 28155 ( SAPS) ; loc. cit., 14 June 1937, Tatewaki 28155 ( SAPS) ; loc. cit., 26 June 2023, Shutoh Da 11 ( KYO) ; Samani-gun, Samani-cho, Fuyushima , Mt. Apoi , 15–19 July 1929, Koizumi 19274 ( TNS) ; loc. cit., date unknown, Tsushima s.n. ( TI) ; loc. cit., 28 June 1974, Kumata s.n. ( SAPS) ; loc. cit., 29 May 1927, Tatewaki 9117 ( SAPS) ; loc. cit., 12 June 1952, Tatewaki & Tsujii s.n. ( SAPS) ; loc. cit., 12 June 1977, Hara 4761d ( SAPS) ; Saru-gun, Hidaka-cho , 5 June 1952, Samejima & Nishimura s.n. ( SAPS) ; Hakodate-shi, Mt. Hakodate , 27 May 1982, Ito 5878 ( SAPS) ; Hokuto-shi, Moheji , 25 May 1907, Ninoue s.n. (SAPS ; Hiyama-gun, Assabu-cho , 24 May 1955, Tatewaki, Igarashi & Watanabe s.n. ( SAPS) ; Nishi-gun, Otobe-cho, Asahitai , Takemori , 16 May 2000, Takahashi 27041 ( SAPS) ; Hiyama-gun, Assabu-cho , 10 May 2012, Suetsugu Da 4 ( KYO) ; loc. cit., 21 October 2021, Sugawahara Da 3 ( KYO). Tohoku District — Aomori Pref.: Mutsu-shi, Ohata-cho , Mt. Osorezan , June 1912, Kinashi s.n. ( KYO) ; Higashitsugaru-gun, Hiranai-cho, Karibasawa , Mt. Eboshi-dake , 10 June 1928, Kudo s.n. ( KYO) ; Mutsu-shi, Kawauchi-cho, Shishihata , Kawauchigawa Valley , 23 May 1996, Koga 12243 ( KYO) ; Nishitsugaru-gun, Hukaura-machi, Juniko , 19 June 1932, Muramatu s.n. ( TI). Iwate Pref.: Kamaishi-shi, Kasshicho, Mt. Goyosan , 8 June 1952, Kawasaki s.n. ( TNS). Akita Pref.: Odate-shi, Hinai-machi, 7 June 1931, Matsuda s.n. ( TNS) ; Oga-shi, Oganakakokuyuchinai, Mt. Kenashi , 30 May 2004, Horii 35073 ( KYO). Fukushima Pref.: Mt. Azuma , July 1903, Nakahara s.n. ( TI) ; Minamiaizu-gun, Hinoemata-mura, Mt. Magohyoeyama , 24 July 1966, Okuda 5370 ( TNS). Kanto District — Tochigi Pref.: Shimotsuke-shi , 20 May 1915, Numajiri s.n. ( KYO) ; Nikkou-shi, Mt. Nyohozan , 20 July 1902, Takeda s.n. ( SAPS) ; Nikkou-shi, near Shizu , 25 July 1902, Takeda s.n. ( SAPS) ; loc. cit., 26 July 1902, Takeda s.n. ( SAPS) ; Nikko-shi, Chugushi, Shizu , 3 July 1931, Maekawa 4300 ( TNS) ; Nikko-shi, Chugushi, Shizu , Mt. Nantai , 7 February 1931, Tsuyama s.n. ( TI) ; Nikko-shi, Yumoto , 16 June 1878, collector unknown s.n. ( TI) ; Nikko-shi, Yumoto, edge of Lake Yunoko , 2 July 1947, Teramoto s.n. ( TI) ; Nikkou-shi, Konsei-ridge, date unknown, Takeda s.n. ( SAPS). Gunma Pref.: Tone-gun, Katashina-mura, Tokura , 19 June 1951, Mizushima 1200 ( TI) ; Tone-gun, Katashina-mura, Tokura , Hatomachi-toge , 5-11 June 1948, Kawasaki 4905 ( TNS) ; loc. cit., 3 July 1941, Hara s.n. ( TI) ; Tone-gun, Katashina-mura, Tokura , north of Mt. Shibutsu , 15 July 1929, Hara s.n. ( TI) ; Agatsuma-gun, Tsumagoi-mura, Tashiro , Mt. Azumaya , 1 July 1934, Hara s.n. ( TI) ; Agatsuma-gun, Nakanojo-cho, Iriyama , 22 June 2023, Matsui, Ichikura & Ichikura Da 13 ( KYO). Saitama Pref.: Chichibu-gun , Yokoze-machi , Mt. Bukozan , 8 June 1941, Shimizu s.n. ( TNS) ; Chichibu-shi, Nakatsugawa Mt. Ryokami , 19 May 1956, Hara s.n. ( TI) ; Chichibu-shi, Nakatsugawa, Jumonji-toge , 8 June 1930, Narita s.n. ( TI). Chubu District — Toyama Pref.: Nakaniikawa-gun , Tateyama-machi , Ashikuraji , Bunazaka , 9 June 1935, Hashimoto s.n. ( TI) ; loc. cit., 7 June 1936, Hashimoto 1384 ( TI). Yamanashi Pref.: Minamitsuru-gun, Nishikatsura-cho, Shimokurechi , Mitsutoge , 10 June 1934, Hiyama 4999 ( TNS) ; Minamitsuru-gun, Narusawa-mura, Fujifunatsuguchi mountain trail, 19 May 1957, Kanai s.n. ( TI) ; Minamitsuru-gun, Fujikawaguchiko-machi, Shoji , Aokigahara , northern foot of Mt. Fuji , 13 June 1965, Kawano, Ohashi & Ihara s.n. ( TI) ; Minamiarupusu-shi, Asuyashiashikura, en route from Momonoki hot spring to Yashagami-toge , 4 June 1967, Konta 6070 ( TNS) ; Hokuto-shi, Kobuchisawa-cho, Mt. Yatsugatake , 9 July 1917, Yamada s.n. ( KYO) ; loc. cit., 18 June 1955, Murata & Horikawa 1076 ( KYO) ; Nirasaki-shi, Maruno-machi, Shimotsuburai , Mt. Hohou , 15 July 1952, Ito s.n. ( TNS). Nagano Pref.: Minamisaku-gun , Kitaaiki-mura , Mt. Ogura , 17 June 1977, Murata 3875 ( TI) ; Kitasaku-gun, Karuizawa-machi , 15 June 1958, Koyama n.21182 ( TNS) ; Minamisaku-gun, Kawakami-mura, en route from Senjogahara to Jumonji-toge , 8 August 1958, Hotta 10225 ( KYO) ; Shimotakai-gun, Yamanouchi-machi, Hirao , Mt. Yakebitai , 13 July 1980, Midorikawa s.n. ( TI) ; Minamisaku-gun, Sakuho-cho, Yachiho Highland , 10 July 1974, Mimoro, Tsugaru & Nishiyama 1846 ( KYO) ; Chino-shi, Minotoguchi, Mt. Yatsugatake , 6 June 1990, Midorikawa 906011 ( TI) ; Chino-shi, Mt. Yatsugatake , July 1923, Oka-Hubo s.n. ( TNS) ; Suwa-gun, Hara-mura, Mt. Akadake , 25 June 1936, Ishizuka 67 ( TI) ; Chiisagata-gun, Nagawa-machi, Daimon , July 1926, Abe s.n. ( TNS) ; Ina-shi, Hasekurogochi, Mt. Senjogatake , 11 July 1965, Iwatsuki, Fukuoka & Naruhashi 197 ( KYO) ; Ina-shi, Hasekurogouchi, Mt. Nyukasa , the point where the Oguro and Minamisawa Rivers converge, 21 June 1954, Kanai s.n. ( TI) ; Shimoina-gun, Oshika-mura, Kashio , Mt. Takamori , 9 June 1957, Asano 10643 ( TI) ; Azumino-shi, Misatoogura, Kurosawa Waterfall , 28 May 1995, Koga 11805 ( KYO) ; Azumino-shi, Hotakaariake, Mt. Tsubakurodake , 25 June 1914, Yazawa s.n. ( TNS) ; Matsumoto-shi, Azumi, Kamikochi , between Okumatashiro and Yokowo, 29 June 1968, Sugiyama 11162 ( TI). Gifu Pref.: Nakatsugawa-shi , Misaka , Fujimidai , 10 June 1965, Asano 9331 ( TI) ; Nakatsugawa-shi, Misaka , 15 May 2022, Iwahori Da 5 ( KYO) ; loc. cit., 5 June 2022, Iwahori Da 6 ( KYO) ; Takayama-shi, Takane-machi, Nomugi , Nomugi-toge , 13 June 1958, Kanai s.n. ( TI) ; Nakatsugawa-shi, Nakatsugawa, Mt. Ena , 19 July 1961, Hashimoto s.n. ( TI) ; Takayama-shi, Nyukawa-cho, Iwaidani , Mt. Norikura , 5 July 1936, Koizumi s.n. ( KYO) ; loc. cit., 2 July 1966, Okuyama s.n. ( TNS) ; Gero-shi, Hagiwara-cho, Sakurabora , Mt. Gozenyama , 23 May 1971, Murata, Okamoto & Takahashi 31 ( KYO). Shizuoka Pref.: Fujinomiya-shi , Awakura , Mt. Fuji , 28 June 2022, Sato Da 8-1 ( KYO) ; Fujinomiya-shi, Mt. Fuji , 1864, Tschonoski (=Sukawa) s.n. (K) ; Fujinomiya-shi, Awakura, Mt. Fuji , 8 June 2022, Sato Da 7 ( KYO) ; Shizuoka-shi, Aoi-ku, Tashiro , Mt. Chausudake , 7 July 1967, Konta, Tsuno & Nagao Konta 6180 ( TNS). Aichi Pref.: Kitashitara-gun , Shitara-cho , Tsugu Honsawa , Mennoki Touge , 18 May 1952, Torii 12439 ( KYO) ; Kitashitara-gun, Shitara-cho, Damine , Dandobenzaiten , 26 May 1951, Torii 12375 ( KYO). Kinki District— Mie Pref.: Mie-gun, Komono-cho, Komono, Mt. Gozaisho , 10 May 1958, Iwatsuki & Kitagawa s.n. ( KYO). Hyogo Pref.: Mikatagun, Kami-cho, Muraokaku Yamada, Mt. Mikawayama , 5 May 1978, Hashimoto 11101 ( KYO) ; Shiso-shi, Chikusacho, Nishigouchi , Chikusa Kogen , 21 May 2000, Kobayashi 34499 ( KYO). Nara Pref.: Yoshino-gun , Tenkawa-mura , Dorogawa , Mt. Sanjogadake , 27 May 1955, Murata & Iwatsuki 98 ( KYO) ; Kamikitayama-mura, Mt. Odaigahara , 22 May 1968, Hotta 16906 ( KYO) ; Yoshino-gun, Tenkawa-mura, Dorogawa , 26 May 1955, Murata & Iwatsuki 532 ( KYO). Wakayama Pref.: Ito-gun , Koya-cho , Mt. Koya , May 1921, Nakajima s.n. ( KYO) ; loc. cit., 1921, Nakajima s.n. ( TI). Shikoku District — Tokushima Pref.: Mima-gun, Tsurugi-cho , Ichiu , Mt. Marusasa , 5 June 1984, Yamazaki s.n. ( TI) ; Miyoshi-shi, Higashiiyasugeoi, Mt.Tsurugi , 5 May 1961, Tabata s.n. ( KYO). Ehime Pref.: Niihama-shi , Besshiyama , Mt. Higashiakaishi , 30 May 1957, Murata & Shimizu 1846 ( KYO) ; Kita-gun, Uchiko-cho, Nakagawa , Odamiyama Valley , 17 May 1925, Yamashita s.n. ( KYO). Kochi Pref.: Agawa-gun , Niyodogawa-cho , Nanokawa , 21 May 1888, unknown collector s.n. (TI0) ; loc. cit., 4 April 1890, Watanabe s.n. ( TI) .

Vernacular name:— Ichi-yo-ran (in Japanese).

Taxonomic notes:— Dactylostalix ringens was described based on specimens from Kii (now Wakayama and southwestern Mie Prefectures) collected in June 1875 by Johannes Justus Rein ( Reichenbach 1878). Given Rein’s travels in June 1875 ( Rein 1881), the most likely location is Mt. Koya, Wakayama Prefecture. The plants currently treated as D. ringens (a common type within the D. ringens species complex) have been found predominantly around this type locality, encompassing not only Mt. Koya but also other areas in the Kinki District, including Wakayama and southwestern Mie Prefectures. Despite the poor preservation of its holotype specimen, line drawings based on this type specimen provide detailed lip morphology, crucial for identifying Dactylostalix species. The semiorbicular lateral lobes align well with typical D. ringens individuals ( Fig. 2 ). Since D. ringens was described as having ligulate and rounded lateral lobes ( Reichenbach 1878), D. ringens was undoubtedly described based on the entity we now recognize as D. ringens .

Calypso japonica was described based on specimens from three localities (Mt. Fuji, Shizuoka Prefecture; Hakone, Kanagawa Prefecture; and Nambu, Iwate Prefecture) collected in 1864–1866 by Sugawa Chonosuke ( Komarov 1901). Reviewing the original materials, we found C. japonica was described based on a mixture of D. ringens and D. uniflora ( Figs. 5–6 ). Specifically, individuals from Nambu (LE 01012252) share characteristics with D. uniflora , such as small, narrowly triangular lateral lobes ( Fig. 5 ). In contrast, most individuals from Mt. Fuji (LE 01012254) align with D. ringens due to large, semiorbicular lateral lobes ( Fig. 6 ). The individual from Hakone (LE 01012253) was indeterminate due to late collection and subsequent difficulty in floral morphology assessment. Given the brevity of Maximovich’s description in Komarov (1901: 533), it is unclear whether D. ringens or D. uniflora was the focus. However, since C. japonica has been synonymized with D. ringens in subsequent studies ( Freudenstein 1994; Freudenstein et al. 2005; Inoue 2016), we designate the D. ringens collected at Mt. Fuji as the lectotype to stabilize the nomenclatural position of C. japonica . Notably, LE 01012254 likely comprises a few individuals of D. uniflora (e.g., the second individual from the left). This supposition is justifiable, as our field surveys confirmed their co-occurrence in Mt. Fuji. To eliminate ambiguity, we have designated a single D. ringens plant (the third individual from the right in the upper row in Fig. 6A ) as the lectotype of C. japonica .

Dactylostalix maculosa was described based on specimens from Moheji, Hokkaido Prefecture, collected on 16 May 1906 and 25 May 1907 by Shimejiro Ninoue. Miyabe & Kudo (1915) noted that D. maculosa differs from D. uniflora by having a broader stem and a larger flower with a roundish labellum featuring conspicuous purple dots. However, these characteristics as outlined by Miyabe and Kudo (1915) align with those of D. ringens . D. maculosa was described based on two specimens collected on different dates. Given that (i) the sheet SAPS036659 was already marked as a type specimen and (ii) the key diagnostic characteristic (large, semiorbicular lateral lobes) was clearly identifiable on the sheet, we designate SAPS036659 as the lectotype and SAPS036660 as the isolectotype of D. maculosa ( Fig. 7 ). The relative prevalence of P. uniflora in Hokkaido likely led to the erroneous assumption by Kudo that D. ringens and P. uniflora were identical, subsequently leading to the description of D. maculosa as a new species.

Dactylostalix ringens f. punctata was described based on a specimen from Kamioboro, Hokkaido Prefecture, collected on 14 June 1937 by Misao Tatewaki. D. ringens f. punctata is distinguished from D. ringens f. ringens solely by the presence of purple-brown spots on the leaves. Since there are no significant differences in labellum and column morphology, which are crucial for identifying Dactylostalix species, we have concluded that D. ringens f. punctata is synonymous with D. ringens . Given that Miyabe & Kudo (1915) had originally named what is now recognized as D. ringens as D. maculosa , it may seem somewhat perplexing that, in Miyabe & Tatewaki (1937), D. ringens f. punctata was correctly identified as a variant of D. ringens . A likely explanation for this discrepancy is that, although Miyabe was the lead author in both publications, it was Kudo and Tatewaki who spearheaded the research and wrote the papers, respectively. It has been suggested that, possibly due to the rules of Hokkaido University at that time, Kudo and Tatewaki often listed Miyabe’s name as the primary author in many papers, a practice contrary to the current norm of including the supervisor as the last author (Shutoh Kohtaro, personal communication). Therefore, it can be inferred that there was a difference in the understanding of D. ringens between Kudo and Tatewaki, with the latter accurately recognizing D. ringens f. punctata as a variant of D. ringens .

Preliminary conservation status: —So far, more than 1000 mature individuals of D. ringens have been observed. This species mainly thrives in densely forested regions, which are often situated within protected areas or forest parks. This species mainly thrives in densely forested regions, which are often situated within protected areas or forest parks. In accordance with the IUCN criteria (IUCN 2019), the conservation status of D. ringens is evaluated as “Least Concern”.

Figures

FIGURE 1. Morphological comparison of Dactylostalix ringens (A–C) and D. uniflora (D–F). (A) Flowering plant in Ena-shi, Gifu Pref. (B–C) Flower in Mt. Fuji, Fujinomiya-shi, Shizuoka Pref. (D) Flowering plant in the Akaishi Mountains, Yamanashi Pref. (E–F) Flower in Mt. Fuji, Fujinomiya-shi, Shizuoka Pref. Scale bars: 30 mm (A, D) and 10 mm (B–C & E–F).

FIGURE 2. Holotype of Dactylostalix ringens (W0196072). (A) Habit and its line drawing. (B) Line drawing of the labellum. (C) Line drawing of the upper part of the column (left) and pollinarium (right). Black arrows indicate semiorbicular side lobes. Scale bars: 50 mm (A) and 5 mm (B–C).

FIGURE 5. Phylogenetic tree of Dactylostalix ringens and D. uniflora reconstructed from MIG-seq data. Nodes supported by bootstrap values <50% are not shown. Branch length represents the average number of substitutions per site.

FIGURE 6. Neighbor-Net network of Dactylostalix ringens and D. uniflora reconstructed based on uncorrected p distance calculated from 6062 SNPs.

FIGURE 7. Original materials of Calypso japonica (LE01012254). (A) Habit. (B) Flower, front view. (C) Flower, lateral view. A black arrow in (A) indicates a lectotype designated here, while black arrows in (B) and (C) indicate semiorbicular side lobes. The individual shown in (B) was designated as a lectotype in the present study. Scale bars: 50 mm (A) and 10 mm (B–C).

FIGURE 9. Lectotype of Dactylostalix maculosa designated in the present study (SAPS036659). (A) Habit. (B) Flower, front view. (C) Line drawing of the flower. Black arrows indicate semiorbicular side lobes. Scale bars: 50 mm (A) and 10 mm (B–C).

FIGURE 10. Holotype of Dactylostalix ringens var. punctatus (SAPS036661). (A) Habit. (B) Flower, front view. (C) Flower, lateral view. Black arrows indicate semiorbicular side lobes. Scale bars: 50 mm (A) and 10 mm (B–C).

FIGURE 11. Dactylostalix ringens (Katsumi Iwahori Da5, KYO). (A) Habit. (B) Flower, lateral view. (C) Column, labellum, and ovary. (D) Dorsal sepal. (E) Lateral petal. (F) Lateral sepal. (G) Labellum. (H) Flatten labellum. (I) Column. (J) Pollinarium on the apical part of the column. (K) Anther cap and pollinarium. Scale bars: 30 mm (A) and 5 mm (B–K).