Spicantopsis niponica (Kunze) Nakai, 1933

Molino, Sonia, Santos, Guillermo, Vázquez, Rubén, Medina, Rafael & Gabriel, José María, 2024, Monograph of the genera Struthiopteris Scop. and Spicantopsis Nakai (Blechnaceae, Polypodiopsida), Phytotaxa 677 (1), pp. 1-48 : 15-17

Spicantopsis niponica (Kunze) Nakai ( Figure 9C ; Figure 10F–G ; Figure 13A, D, E ).

Type:— JAPAN. Hondo, Suwo, Hikami oppida Ouchi, no date, J. Nikai 12 ( TI 00274317! photo, neotype, designated here).

Synonyms:— ≡ Lomaria niponica Kunze. Blechnum niponicum (Kunze) Makino (1897: 82) . Blechnum spicant ( L.) Roth (1794: 56) var. niponicum (Kunze) Christensen (1906: 405) . Spicanta niponica (Kunze) Hayata (1927: 700) (original: ‘ nipponica ’). Struthiopteris niponica (Kunze) Nakai (1930: 15) (original: ‘ nipponica ’).

= Lomaria spicant (L.) Desvaux (1811: 350) var. japonica Hooker (1860: 15) (original: ‘ β. Japonica ’). Type:— JAPAN. Hakodadi, 1853‒ 56, C. Wright s.n., North Pacific Exploring Expedition ( US bc01527323!, holotype) ≡ Spicantopsis niponica (Kunze) Nakai (1933: 181) var. japonica (Hook.) Nakai (1933: 181) .

= Blechnum spicant (L.) Roth var. reflexipinnula Masamune (1932: 303) . Type:— JAPAN. Yakushima, 1928, Aikodake (not found, synonymy based on the protologue and current usage of the name). ≡ Spicantopsis niponica (Kunze) Nakai var. reflexipinnula (Masam.) Nemoto (1936: 75) . Struthiopteris niponica (Kunze) Nakai var. reflexipinnula (Masam.) Masamune (1951: 198) .

= Blechnum spicant (L.) Roth var. undulatipinnula Masamune (1932: 303) . Type:— JAPAN. Mt. Takakuma , 1922, no collector (not found, synonymy based on the protologue and current usage of the name) ; ≡ Spicantopsis niponica (Kunze) Nakai var. undulatipinnula (Masam.) Nemoto (1936: 75) .

= Spicantopsis niponica (Kunze) Nakai f. flabellata Satô (1936: 824, Fig. 2 ). Type:— JAPAN. Insula Yakushima, inter Kosugidani et Nakano-Koya, 24 August 1935, K Satô s.n. (not found, synonymy based on the protologue and current usage of the name). ≡ Struthiopteris niponica (Kunze) Nakai (1933: 181) f. flabellata (Satô) Namegata & Kurata (1961: 332) .

= Struthiopteris niponica (Kunze) Nakai f. kinoshitae Itô (1971: 38) . Type:— JAPAN. Shikoku: mamidani, Tazukawa, Katsuura-machi, Katsuura-gun, Tokushima Pref., 10 October 1970, S. Kinoshita s.n. (TI00081476! photo, holotype).

Plants terrestrial; rhizomes thick, short and erect, covered by linear scales (0.6) 1‒2 x (2) 11‒16 mm, papyraceous, discoloured, centre broadly sclerotized from base to apex, red or dark brown, margins narrow, light brown, entire; fronds dimorphic, sterile fronds (15) 33 (51) cm long, short petiolate, stipes light brown or green, 0.5‒6 cm, laminae linear to elliptic-lanceolate, coriaceous, (3) 5‒8 (10) x 15‒30 (48) cm, 1-pinnate, pinnatifid or pinnatisect, apex conform, abaxially with small scales, brown, triangular-lanceolate; rachises grooved adaxially, abaxially with scales similar to those on the lamina, pinnae tapering towards the base, the middle ones 0.2‒0.7 x 1‒3 (7) cm, costae grooved adaxially, prominent abaxially, veins free, simple or 1-furcate, catadromous, ending in submarginal patent hydathodes; fertile fronds distinctly longer than sterile ones, (19) 40 (67) cm long, stipes light brown or green, 2‒5 (14) cm, laminae linear to elliptic-lanceolate, 13‒35 (54) cm, with scales on the abaxial side of the lamina and rachis like those of the sterile fronds; pinnae contracted, the middle ones 0.1‒0.4 x 1.5‒3 (5) cm; sori linear, continuous, forming coenosori on both sides of costa, occupying entire length of pinna, not decurrent towards rachis; indusia brown, continuous, long and coiled; sporangia with 19‒20 arc cells; spores monolete, 30‒55 x 50‒70 µm, perisporium brown, surface granularly reticulate.

Habitat and distribution:— Endemic to Japan. Throughout Japan, from Hokkaido to Yakushima (Kyushu) ( Figure 7 ) ( Iwatsuki 1992, Itwasuki et al. 1995). On village roadsides and cleared fields at the foot of hills ( Iwatsuki 1992) as well as in montane forests ( Ohwi 1953). 60‒1600 m.

Chromosome number:— 2 n = 62, diploid ( Fabbri 1965, Nakato 1987).

Etymology:— The epithet niponica refers to Japan, where the plant is native. Sometimes the specific epithet is found as nipponica . This spelling is not correct as both in the original publication of Spicantopsis niponica ( Nakai 1933) and in the basionym Lomaria niponica ( Kunze 1848: 508) is found with a single p.

Vernacular name:— Shishigashira (シシガシラ, Japanese) ( Iwatsuki 1992).The vernacular name “shishigashira” means “lion head”, probably referring to the rhizome scales, which remain of male lion hair.

Taxonomic notes:— This species closely resembles Spicantopsis hancockii , however it differs in some aspects (see section on S. hancockii ), of which the most reliable seems to be the presence of two posterior cells in the sporangium of S. niponica , absent in S. hancockii . Moreover, recent studies have shown that these two species do not coexist, S. niponica being a Japanese endemic and S. hancockii a Taiwanese endemic. It is also similar to its congener S. amabilis , but the size of the plant and differences in the rhizome scales easily separate them (see S. amabilis section).

Several forms have been recognized in Yakushima (see synonyms section), where its other variety, Spicantopsis niponica var. minima , was also described, which shows that it is a species with high phenotypic variability ( Iwatsuki 1992). The niponica variety can be easily distinguished from the minima variety by its size, with S. niponica var. minima not exceeding 15 cm.

The original material cited in Kunze (1848) is: Japonia. Göring s.n., no date (Herb. Klenz. Lucaean. et proprium). Kunze’s herbarium was deposited in Leipzig (LZ) and mostly destroyed during the Second World War, as confirmed by LZ staff and as stated in Thiers (2024. Continuously updated), making the designation of a neotype necessary. In the absence of a specific locus classicus, we have selected one of the specimens cited by Nakai in his combination Spicantopsis niponica ( Nakai 1933) as a neotype: J. Nikai 12 (TI00274317).

Additional specimens examined:— JAPAN. Hokkaido. no date, no collector s.n. ( TI00274319 ! photo) ; no date, N. C. Wright s.n. ( US bc01527323!). Pref. Hokkaido: forest reservation near Sapporo , September 1929, P. H. Dorsett & W. J. Morse 1038 ( NL3527438 ! photo; US1577946 !) ; Ishikari, Sapporo , illegible date, R. Yatabe ( TI00274318 ! photo) ; Ishikari, Sônkyôu , June 1879, T. Nakai s.n. ( TI00274320 ! photo) ; Mt. Yotei, lake Hangetsu , 2 August 1945, M. Mizushima s.n. ( TI00274329 ! photo) ; Ori river, Homme, Shimamaki-mura , Shimamaki-gun , Shiriboshi-shicho , 3 August 1980, M. Kato & D. E. Boufford 1581 ( GA088401 ! photo) ; Oshima , 25 July 1931, illegible collector 222 ( TI00274262 !photo) ; Oshima, Hakodate , 4August 2008, Y.- M. Huang s.n. ( TAIF297604 ! photo) ; Ibidem ( TAIF297605 ! photo) ; Sapporo , 12 October 1903, S. Arimoto s.n. ( MO1877400 !) ; Ibidem , September 1929, P. H. Dorsett & W. J. Morse 1038 ( P01575938 !). Honshu. 1866–1871, Kramer 2280 ( P01406581 !) ; 1866, Tschonoski s.n. ( P01406591 !) ; Ibidem ( P01406593 !). Pref. Aichi: Kitashidara-gun , Shidara-machi , 13 August 1977, J. Murata 4106 ( TI00274391 ! photo) ; Omoteyama, 7 August 1931, T. Nakai s.n. ( TI00274314 ! photo). Pref. Akita: Betsudake , Nishikurosawa , 24 June 1946, T. Yamazaki s.n. ( TI00274349 ! photo) ; Okachi-gun, Okachi-machi, Matsunoki pass, 8 October 1972, H. Kanai & Y. Tateishi 1110 ( TI00272835 ! photo; US2740562 !). Pref. Aomori: 1866, J. Faurié 146 ( P01406584 !) ; Agamushi , 12 November 1912, J. Faurié 12 ( NY04179851 !) ; Ibidem , 12 November 1913, J. Faurié ( P01406533 !) ; Aomori-ken, Bonjusan northwest of Daishaka , 24 August 1928, R. K. Beattie & Y. Kurihara 10212a ( US2292145 !) ; Ibidem , R. K. Beattie & Y. Kurihara 10312 ( US2292144 !) ; Hakkoda, no date, no collector s.n. ( TI00274345 ! photo) ; Kamikita Dist., Towada , 2 November 1946, U. Narita 555 ( TI00274339 ! photo) ; Mutsu, Mt. Hakkoda , 31 July 1926, S. Hattori s.n. ( TI00274316 ! photo) ; Towada, 26 June 1894, E. Drake & J. Faurié 13280 ( C0364773 F!; C0364775 F!; P01406585 !). Pref. Fukui: Echizen, Nanjo Nagono, 26 July 1933, Y. Hosoi 151 [interpreted] ( TI00272857 ! photo). Pref. Chiba: Tsui, Komachi, Shimosa, Tsuba, Ichikawa, Matsu, 14 July 1963, H. Ohba 418 ( TI00274400 ! photo). Pref. Fukushima: Aizu, the foot of Mt. Bandai-san , 3–4 November 1967, H. Ohba 671116 ( TI00274384 ! photo) ; Hanami Yama , 1866–1871, L. Savatier s.n. ( P01406583 !) ; Haramachi-shi , 16 December 2004, K. Iga 24 ( TI00203872 ! photo) ; hills north of Hamata , 24 August 1927, R. K. & Y. Kurihara 10047 ( US2292107 !) ; Iwashiro , 14 July 1940, T. Nakai s.n. ( TI00274279 ! photo) ; Iwashiro, Adizu , August 1879, R. Yatabe & J. Matsumura s.n. ( TI00274323 ! photo) ; Kaihozan , 1 October 1949, H. Uematu 1780 ( TI00274360 ! photo) ; Nishishirakawa, Furomuraya , 24 May 1930, H. Sakaki 54 ( TI00274269 ! photo) ; Yama-gun, Yamato-machi, Ichinoki , 28 August 1980, H. Ohba & S. Akiyama 1867 ( TI00274294 ! photo). Pref. Gifu: Mt. Kinka , 1 November 2010, W.- L. Chiou 15469 ( TAIF462180 ! photo; TAIF462184 ! photo) ; Ono, Miwa Sanctuary , 8 June 1886, S. Tanaka s.n. ( TI00272845 ! photo). Pref. Gunma: Kôdzuke, circa cataractum Ogura, Shima, August 1929, T. Kondô s.n. ( TI00274330 ! photo) ; Kusatsu , September 1877, Dickim [interpreted] ( P01406587 !) ; Mt. Akagi , November 1934, S. Adachi 28 ( TI00272855 ! photo) ; Takasaki-shi, Mt. Kan’nonyama , 23 December 1966, H. Ohba 662135 ( TI00274385 ! photo) ; Tano-gun, Ueno-mura, Otchizawa , 15 July 1976, J. Murata 2252 ( TI00272859 ! photo) ; Tano-gun, Ueno-mura, Yaegoya , Jukkokutooge , 8 September 1976, J. Murata 2824 ( TI00272860 ! photos). Pref. Hiroshima: Saiki-gun, Itsukaichi, Mt. Madogayama , Ugiri valley , April 1971, T. Koponen & A. Koponen 16808 ( H1675159 ! photo) ; Ibidem , T. Koponen & A. Koponen 16817 ( H1675149 ! photo) ; Tentokukyo, 1936, illegible collector ( TI00274280 ! photo). Pref. Hyogo: Awaji , Toda , 6 August 1948, A. Ishigami s.n. ( TI00274369 ! photo) ; Hikami-gun, Mt. Myoko-san , 23 September 1960, M. Tagawa & K. Iwatsuki 3014 ( TI00274394 ! photo) ; Kawakami, Oogochicho, Kanzaki-gun , 26 July 1965, K. Iwatsuki 6988 ( US2424173 !) ; Mt. Myôkô-san, Hikami-gun , 23 September 1960, M. Tagawa & K. Iwatsuki 3014 ( P01575937 !; US2423733 !) ; Yamamoto in Sottsu , 10 February 1953, N. Togasi 710 ( TI00274366 ! photo) ; Yamamoto, Takarazuka, 28 July 1962, M. Togashi s.n. ( TI00274395 ! photo). Pref. Ishikawa: Keta Sanctuary , 29 June 1952, N. Satomi s.n. ( TI00272853 ! photo) ; Komatsu-shi, Sawa-machi , 22 August 1994, K. Yonekura 3331 ( MO04989202 !) ; Mt. Sekido , 26 June 1972, M. T. Kao 8405 ( NY04179848 !) ; Noto, Minani , 11 July 1931, H. Hara s.n. ( TI00274258 ! photo) ; Noto, Ushitsu, Yaho , 13 February 1931, H. Hara 95 ( TI00274256 ! photo). Pref. Iwate: Nishiiwai Gun, Hiraizumi Machi, Motsuji Temple, 1 November 2018, W.- C. Hsu 72 ( TAIF522703 ! photo). Pref. Kanagawa: 25 October 1889, no collector ( TI00274374 ! photo) ; Hakone , July 1869, Drake 1558 ( P01406579 !) ; Hakone , 1866‒1974, M. Savatier 1558 ( US483272 !; US bc05127319!) ; Hakone, Atami michi, 26 December 1886, no collector s.n. ( TI00274284 ! photo) ; Hakone, Komagatake , 1 August 1906, H. Sakurai s.n. ( P01406576 !) ; Hakone, Sagami , 21 October 1902, M. Weigel s.n. ( P01406578 !) ; ibidem, August 1904, T. Makino s.n. ( TI00274285 ! photo) ; Kantō, Yokohama, 1862, Maximowicz s.n. ( NY04179855 !) ; Ibidem ( P01406592 !) ; Sagami , 24 July 1962, H. Ohba 10911 ( TI00274299 ! photo) ; Sagami, Tanzawa , 22 July 1942, illegible collector 10814 ( TI00274298 ! photo). Pref. Kyoto: 1891‒1893, K. Berry s.n. ( MO2817713 !) ; Canyon on Otani-yama back of Oji, Tamba , 2 November 1927, R. K. Beattie & Y. Kurihara 10128 ( US2292118 !) ; Chohroh-ga-take ( Mount Chorogatake ), 9 September 1979, Y. Saiki 1022 ( C0364727 F!) ; Fushimi Inari Taisha Shrine , 6 October 2015, C.- W. Chen Wade 4529 ( TAIF500147 ! photo) ; Hozu-chō, Kameoka-shi , 14 September 1987, T. Takahashi 423 ( MO3893847 !) ; Kitakuwada, Chii, Experimental Forest of Ashio University , 17 July 1940, T. Nakai s.n. ( TI00274277 ! photo) ; Kurama , November 1895, no collector ( NY04179857 !) ; Miyama-cho, Ashiu, Kyoto University Forest Reserve , 19 September 1996, H. van der Werff 14052 ( MO0536479 !) ; mountain back of Miyako Hotel , 25 May 1929, R. K. Beattie & Y. Kurihara ( US2292160 !) ; Ohmi, Mt. Hiei , 11 September 1966, van Steenis s.n. ( NL3527437 ! photo) ; Sakura-toge, Higashibetsuin-cho, Kameoka-shi , 16 August 1991, T. Takahashi 2060 ( MO4029368 !) ; Sakyo-fu, tributary of Kitashiro-gawa river, SW side of Mt. Hiei-zan , 19 June 1977, D. E. Boufford & E. W. Wood 19188 ( NL984166232 ! photo) ; Shishigatani , 30 October 1955, M. Tagawa 7150 ( P01608625 ; TI00274305 ! photo; US2420041 !) ; Tango , August 1905, K. T. Tanaka s.n. ( TI00274292 ! photo) ; Yokosuka, 1866–1871, A. Henschel 1558 ( P01557335 !). Pref. Mia: Iga , Ishiharadani , 8 august 1880, J. Matsumura s.n. ( TI00272842 ! photo) ; Ise, Mt. Tarusakayama , 24 August 1920, K. Murata 41 ( TI00272849 ! photo) ; Kitamuro-gun, Yamami-machi, near Kozu-gawa river , 26 July 1981, S. Akiyama 481 ( TI00274306 ! photo) ; Mita Zinsya, north of Ueno , 2 August 1954, M. Tagawa 5928 ( US2259007 !). Pref. Miyagi: Kinkasan, 20 August 1965, H. Ohba 1728 ( TI00272834 ! photo) ; Momou-gun, Okachi-machi, Togezahi , 7 February 1989, T. Kawahara 5606 ( TI00274226 ! photo) ; Rikuzen, Mt. Funagata , 22 November 1974, M. Takahashi s.n. ( MO2321470 !) ; ibidem, 9 July 1972, H. S. Ogura 3538 ( TI00274265 ! photo) ; Rikuzen, Mt. Kattadake , 16 August 1898, Y. Yabe s.n. ( TI00274344 ! photo) ; Sendai-shi, Toshogu , 19 September 1915, Y. Ogura s.n. ( TI00274393 ! photo) ; Tamatsukuri-gun, Narugo-machi, Onikobe Spa , 28 August 1970, H. Ohba et al 708215 ( MO2373914 !). Pref. Nagano: Kiso-gun , Ookuwa-mura , 14 August 1977, J. Murata 4156 ( TI00274392 ! photo) ; Nakamachi , 29 September 1950, K. Sato 380 ( TI00274378 ! photos) ; Nishichikuma-gun, Ookuwa-mura, along the river Aderagawa , 14 June 1979, J. Murata 7720 ( TI00274301 ! photo) ; Nishikoma , 23 June 1972, M. T. Kao 8315 ( NL996175225 ! photo; NY04179847 !) ; Oomachi-shi, Taira, West of Lake Aoki , 20 August 1979, H. Ohba & S. Akiyama 543 ( TI00274387 ! photo) ; Shinano, Minamisaku-gun , June 1938, K. Sato 1419 ( TI00274377 ! photo) ; Shinano, Shimoina, Kazekoshi , August 1940, J. Furusaura s.n. [interpreted] ( TI00274397 ! photo) ; Shinnoa, Kiso, Mt. Ontake , August 1910, G. Koidzumi s.n. ( TI00274273 ! photo) ; Shinnoa, Mt. Yarigatake , July 1920, T. Nakai s.n. ( TI00274254 ! photo) ; Shinnoa, pede montis Yatsugatake , no date, T. Kondô 28 ( TI00272850 ! photo) ; Shinnoa, Wadatôge , 24 July 1880, J. Matsumura s.n. ( TI00274282 ! photo) ; Shinnoa, Zenkôji , 9 July 1884, J. Matsumura s.n. ( TI00274355 ! photo) ; Shinshu, no date, Kondo 28 ( TI00274272 ! photo) ; ibidem, August 1933, H. Isu s.n. ( TI00274276 ! photo) ; ibidem, 14 August 1933, S. Momose s.n. ( TI00272848 ! photo) ; ibidem ( TI00272852 !photo) ; ibidem, 15August 1933, S. Momose s.n. ( TI00272851 ! photo) ; ibidem, November 1933, Ikane [interpreted] ( TI00274281 ! photo) ; ibidem, 25 July 1939, S. Gomi 4 [interpreted] ( TI00272856 ! photo) ; Shinshu, Ushibo, Onsen , 19 January 1933, S. Momose s.n. ( TI00272847 ! photo). Pref. Nara: 13 September 1993, И.М.Красноборов & M. H. Ломоносова s.n. ( NS0009500 ! photo) ; 27 May 1929, P. H. Dorsett & W. J. Morse 711 ( US1577934 !) ; gorge of the Sannoko, Kawakami-mura, Yoshino-gun , 7 October 1954, M. Tagawa 6701 ( US2259054 !) ; Kamikitayama-mura, Yoshino-gun, en route from Mt. Wasamata to Mt. Daifugen , 21 September 1986, S. Tsugaru et al 7443 ( MO4012484 !; NY4179846 !) ; Mt. Kasuga , October 1921, T. Makino 41289 ( TI00274286 ! photo). Pref. Niigata: Minamikanbara-gun, Shimota-mura, near Nagasawa, 17 August 1980, H. Ohba & S. Akiyama 1746 ( TI00274296 ! photo) ; Nakasatomura , 19 September 1982, T. Takahashi 5538-3567,6644 ( TI00138456 ! photo) ; Niitsu-shi , 11 October 1982, T. Takahashi 5639-4089, 6726 ( TI00139368 ! photo) ; Oguni-achi, 11 September 1983, T. Takahashi 5538-7563, 7012 ( TI00221055 ! photo). Pref. Okayama: 14 October 1925, no collector 10220 ( TI00274275 ! photo). Pref Okayama: Maniwa, Kawakami, 2 June 1930, Secondare School of Tokuda s.n. ( TI00274333 ! photo). Pref. Osaka: Yamamoto, Sottsu, 10 February 1953, N. Togashi 710 ( S2061 !). Pref. Saitama: beside Akasawa Valley , the deepest branch of Arakawa River , Kawamata, Ohtaki-mura, Chichibu-gun, 1 August 1963, H. Ohba 535 ( TI00274295 ! photo) ; between Kabagoya and Karisaka-pass, Mts. Chichibu , Ohtaki-mura , Chichibu-gun , 4 August 1963, H. Ohba 562 ( TI00272833 ! photo) ; Kamikawa Division, Asahikawa City, Kaguraoka , 25 July 1924, S. Saito s.n. ( TI00274381 ! photo) ; Mt. Hidaka , August 1928, T. Nakai s.n. ( TI00274309 ! photo) ; Musashi, Mt. Bukôsan , 31 March 1908, T. Nakai s.n. ( TI00274315 ! photo). Pref. Shiga: Ueno Tsuchiyama (Moji), 24 April 1942, T. Hashimoto 286 ( TI00272836 ! photo). Pref. Shimane: Izumo, 5 November 1904, K. Hirata 34 ( TI00274268 ! photo) ; Mt. Daimanji, Oki Islands, 23 July 1932, Y. Nozu s.n. ( TI00274379 ! photo). Pref. Shizuoka: Izu , Kamo Dist. , Minaitsuchi , 28 August 1943, Y. Kimura s.n. ( TI00274365 ! photo) ; Izu lake, Yugashima ( Mukai , Taira ) 11 Nonvember 1946, T. Harada 843 ( TI00274335 ! photo) ; Izu, Mikurajima , October 1932, Takashi s.n. ( TI00274367 ! photo) ; Mt. Fuji , 1924–1926, B. Hayata s.n. ( TI00272840 ! photo) ; Prov. Totomi, the side of Mt. Hakkozan, Oshiro National Forest , Haibara-gun , 16 September 1967, H. Obha 3553 ( TI00274388 ! photo; TI00274389 ! Photo; TI00274390 ! photo) ; Toi River, Toi-cho, Kameda-gun , Oshima-shicho , 26 July 1980, M. Kato & D. E. Boufford 1978 ( GA088400 ! photo) ; Totomi, Nishikashima, Tenryu-shi , 17 June 1961, K. Satake s.n. ( US2979919 !) ; Umegashima-mura, Abetoge, 3 August 1959, Y. Asai s.n. ( TI00274288 ! photo). Pref. Tochigi: Mt. Nasu Daimaru hot spring Shimotsuke, 15 July 1953, T. Namegata 3149 ( US2138707 !) ; Nikko, no date, Ito s.n. ( TI00274352 ! photo) ; ibidem ( TI00274353 ! photo) ; Nikko , 10 September 1808, Y. Numajiri s.n. ( TI00274372 ! photo) ; Nikko , July 1937, J. Furusawa s.n. [interpreted] ( TI00274371 ! photo) ; Shimotsuke, Mt. Nasusan , 12 August 1908, W. Sakourai s.n. ( P01406595 !) ; Ibidem , 16 August 1911, no collector ( US1095352 !) ; Shimotsuke , 3 July 1933, H. Ito s.n. ( TI00274308 ! photo) ; Shimotsuke, Nikko , 1 August 1877, J. Matsumura s.n. ( TI00274327 ! photo) ; Shimotsuke, Shimotsuga-gun, Kawakayama , 2 October 1938, T. Maekawa 80222 ( TI00274368 ! photo). Pref. Tokyo: Bushu , Tama village , 12 August 1946, M. Mizushima s.n. ( TI00274337 ! photo) ; Izu Islands, Island Hachijo-jima, in the crater basin of Mt. Hachijo-fuji Hachijo-cho , 6 November 1977, M. Kato & E. Miki 43 ( TI00274264 ! photo) ; Izu Islands, Island Mikura-jima, July-August 1967, H. Ohba 677185 ( TI00274293 ! photo; US2598946 !) ; Ibidem , 28 July-4 August 1967, Y. Jotani et al. s.n. ( TI00274399 ! photo) ; Izu Islands, Island Miyake , 14–16 April 1983, N. Murakami s.n. ( TI00272861 ! photo) ; Island Nijima, Mt. Mukai , 12 June 2004, T. Miyazaki N4050 ( TI00274382 ! photo) ; Nishitama, Mt. Atago , 10 November 1950, M. Mizushima s.n. ( TI00274270 ! photo) ; Oyama, Miyakejima, 26 August 1979, no collector ( TI00274267 ! photo). Pref. Tottori: Hinogun, Nichinan-machi, Nr. Shimoabire, 9 September 1980, H. Ohba & S. Akiyama 2191 ( TI00274297 ! photo) ; Tottorishi, 17 July 2017, B. Estébanez s.n. ( MACB110656 !). Pref. Toyama: Mugiton , October 1885, Y. Etsu s.n. ( TI00272858 ! photo) ; Neigun, Yatsuo Machi, Tochiori , 12 October 1990, J. Jutila et al. 239 ( MO4380674 !) ; Takano, Tosa, Takaoka , illegible date, K. Yana s.n. ( TI00274336 ! photo). Pref. Yamagata: 23 July 1942, Yamazaki s.n. ( TI00274347 ! photo) ; Mikawa, Okazaki , 5 October 1911, J. Umemura 37 ( P01406521 !) ; Nakayama, Miki-cho, Kida-gun , Kagawa , 5 August 1957, H. Kawasaki s.n. ( TI00274340 ! photo). Pref. Yamaguchi: Kika Park, East slope of Castle Mountain (Shiroyama) National forest , west bank of Nishiki River , Yokoyama village in Iwakuni City. (25.6 miles west of Hiroshima City ), 4 October 1953, L. A. Charette 1441 ( US2256115 !; US2357200 !). Pref. Yamanashi: Kaikoma, 16 July 1946, H. Nematsu s.n. ( TI00274362 ! photo) ; Kai, Mitsutoge , 29 August 1936, K. Hiyama 7760 ( TI00274338 ! photo) ; Kai, Nanri, March 1949, H. Nematsu s.n. ( TI00274361 ! photo). Pref. Wakayama: Hidaka Dist. , August 1928, T. Nakai s.n. ( TI00274311 ! photo) ; ibidem ( TI00274312 ! photo) ; ibidem ( TI00274313 ! photo) ; Hidaka Dist., Samukawa , August 1920, no collector s.n. ( TI00274271 ! photo) ; Kii, Nachi , June 1926, T. Nakai s.n. ( TI00272838 ! photo; TI00272839 ! photo; TI00274255 ! photo; TI00274357 ! photo) ; Kii, Wakanoura , 5 January 1878, J. Matsumura s.n. ( TI00272843 ! photo) ; Koya-san , 23 June 1929, R. K. Beattie & Y. Kurihara 10835 ( US2292165 !) ; Tanabe-shi, Nakahechi-cho, Mizukami , 21 August 2017, B. Estébanez s.n. ( MACB110657 !). Kyushu. North Kyūshū , September 1933, R. K. Smith 20 ( US1756269 !). Pref. Fukuoka: Kokura, August 1929, T. Kondo s.n. ( TI00274331 ! photo) ; Nanokawa, Tosa, 29 July 1892, no collector ( US bc01527320!). Pref. Kagoshima: Amagi , 28 December 1890, no collector ( P01406575 !) ; Amagi , 21 October 1944, Y. Yamazaki s.n. ( TI00274348 ! photo) ; Amagi , 9 October 1956, Owi & Okamoto 1402 ( P01406594 !) ; Amagi, MT. Douzhou , 10 June 1883, no collector s.n. ( TI00272841 ! photo) ; Hanamaki, Inaodake , 14 July 1967, Y. Ando s.n. ( TI00274398 ! photo) ; Kirishima , 10 August 1886, S. Tanaka s.n. ( TI00272844 ! photo) ; Kirishima, 10 August 1938, Department of Agriculture Sciences 4 ( TI00272837 ! photo) ; Mt. Amagi in Idzu , 9 October 1956, J. Ohwi & K. Okamoto 1402 ( H1127512 ! photo; NY04179849 !; TI00274263 ! photo; US2259137 !) ; Ôsumi, Mt. Kirishima , 5 August 1882, R. Yatabe & J. Matsumura s.n. ( TI00274321 ! photo) ; ibidem, 29 August 1890, S. Tanaka s.n. ( TI00272846 ! photo) ; Yakushima Island , 16 July 1922, G. Masamune s.n. ( NY04179853 !) ; Ibidem , 23 September 1984, D. Sather & G. Caddell 6441 ( NCU00348585 ! photo) ; Yakushima Island, around Sandaisugi and Kosugidani , 17 July 1984, M. Kato et al. 17-19 ( TI00274307 ! photo) ; Yakushima Island, Hananoegoo , 6 August 1961, M. Furuse 38848 ( TI00274304 ! photo) ; Yakushima Island, Prov. Ohsumi , 27 December 1961, S. Kurata 6110 ( US2979912 !) ; ibidem 29 July 1924, G. Masamune ( TI00274375 ! photo) ; Yakushima Island, Sandaisugi, Kosugidani Valley , 30 March 1966, H. Ohba 662558 ( TI00274386 ! photo) ; Yakushima Island, Yaku-cho, Hana-yama , Okawa-rindo , 12 October 1977, D. E. Boufford 20171 ( MO2609648 !). Pref. Kanagawa: Sagami, Hakone, August1926, T.Nakai s.n. ( TI00274253 ! photo). Pref. Kumamoto: Fukurengi, Tsunoyama, 3 April 1932, H. Hara 437 ( TI00274257 !photo) ; Hondo, Yamamoto in Sottsu , 10 February 1953, H. Togasi 710 ( H1127514 ! photo; H1127515 ! photo; US2136494 !) ; Mizukami , 19 October 1928, H. Hara s.n. ( TI00274261 ! photo). Pref. Nagasaki: 1863, Maximowicz s.n. ( S2144 !) ; Hizen , May 1879, R. Yatabe & J. Matsumura s.n. ( TI00274356 ! photo) ; Hizen, Mt. Unzen , 6 July 1913, B. Hayata s.n. ( TI00274341 ! photo) ; Unzen National Park, growing on Mount Kunimidake, 28 September 1983, D. Sather & G. Caddell 3026 ( NCU00348584 ! photo). Pref. Ooita: Aso National Park , growing on Mount Kuju along trail from Chojobaru, 9 May 1983, D. Sather & G. Caddell 2517 ( NCU00348583 !photo) ; Takeda-shi, Shirouzu, Ooshoojiiwa ( Mts. Sobo-Katamuki ), 7 July 1979, J. Murata 7979 ( TI00274302 ! photo). Pref. Saga: Mt. Kurokami , August 1950, T. Hashimoto 790 ( TI00274334 ! photo) ; Shikoku. Pref. Ehime: Iyo Prov., Mr. Onigajyô , 8 June 1927, T. Nakai s.n. ( TI00274252 ! photo; TI00274358 ! photo; TI00274359 ! photo). Pref. Kouchi: Hata Distr. , Kuroshio Town , Tachibana-gawa, Kami- Nadayama national forest, 26 October 2010, T. Miyazaki 1011857 ( TI00274364 ! photo) ; Nagaoka-gun, Ootoyo-machi, Mt. Kajigamori , 15 August 1961, K. Oonishi 289344 ( TI00274363 ! photo) ; Takaoka District, Naro, Kubokawa Town , 10 August 2005, T. Miyazaki 508123 ( TI00274290 ! photo) ; Takaoka District, Natsugare Pass, Nakatosa Town , 7 August 2005, T. Miyazaki 508017 ( TI00274289 ! photo) ; Takaoka District, Sakawa-Cho , 19 September 1985, N. Inagaki M85-237 ( TI00274287 ! photo) ; Takaoka District, Shimanto Town, Shiwamine , 1 November 2010, T. Miyazaki 1011239 ( TI00274291 ! photo) ; Takaoka District, Shuujin, Kubokawa Town , 10 August 2005, T. Miyazaki 508146 ( TI00274383 ! photo) ; Tosa , July 1889, Christ s.n. ( P01406589 !) ; Tosa, Mt. Ushioe , 4 January 1939, H. Itô s.n. ( TI00274350 ! photo) ; Tosa, Tosa-gum, Tosayama-mura , 8 June 1931, H. Itô s.n. ( TI00274278 ) ; Ysgi, Asagun, Aki , 8 August 1932, T. Tsuyama ( TI00274396 ! photo). Pref. Tokushima: Awa , Mt. Ôyama , no date, J. Nikai 2320 ( TI00274354 ! photo) ; Mt. Tsurugisan, Nishijimajinja-Taikenjinja , 4 October 1978, J. Murata 6567 ( TI00274300 ! photo) ; Awa, oppido Shimomoyô , 25 July 1888, R. Yatabe ( TI00274322 ! photo). Unknown location: no date, no collector ( P01406582 !) ; ibidem ( TI00274370 ! photo) ; ibidem ( TI00274346 ! photo) ; no date, no collector 345 ( TI00274343 ! photo) ; illegible date, no collector s.n. ( TI00274324 ! photo) ; ibidem ( TI00274325 ! photo) ; ibidem ( TI00274326 ! photo) ; ibidem ( TI00274328 ! photo) ; no date, A. W. Stanford s.n. ( C825009 F!) ; Ibidem ( NY04179856 !) ; no date, Y. Yoshii s.n. [interpreted] ( TI00272854 ! photo) ; 1866‒1871, illegible collector ( P01619264 !) ; 1866, Tschonoski s.n. ( NY04179852 !) ; 1888, H. Matzumo 12998 [interpreted] ( P01619263 !) ; May 1888, no collector ( US bc01527324!) ; June 1896, M. L. Holbrook 177 ( NY04179854 !) ; 8 October 1899, Baker 178 ( MO1877401 !) ; 25 August 1905, H. Sakurai 1912 ( H1127513 ! photo) ; 20 August 1913, no collector ( P01406574 !) ; 18 September 1921, no collector ( TAIF50586 ! photo) ; no date, J. Matsumura 48 ( P01619262 !) ; 12 August 1908, L. Sakurai s.n. ( NY04179850 !) ; 9 February 1910, Christ s.n. ( P01406588 !) ; 12 August 1908, Nerville s.n. ( P01571953 !) ; August 1928, T. Nakai ( TI00274310 ! photo) ; 22 July 1931, H. Muramatsu s.n. ( TI00274274 ! photo) ; Nyokoji [interpreted], 6 August 1934, K. Ohki s.n. ( TI00274332 ! photo) ; 31 January 1937, I. Simozawa s.n. ( MO5423293 !) .

Figures

FIGURE 2. Distribution of Struthiopteris spicant extracted from GBIF. Purple circles: Struhtiopteris spicant var. spicant; yellow squares: Stuthiopteris spicant var. pradae (added manually); blue pentagons: Struthiopteris spicant var. homophyllum (added manually).

FIGURE 7. Distribution of Spicantopsis niponica extracted from GBIF. Purple circles: S. niponica var. niponica; yellow squares: S. niponica var. mínima (added manually).

FIGURE 9. Spores of Spicantopsis: A. S. amabilis (J. Faurié, 5612 P01406648). B. S. hancockii (R. Knapp 4311, P02435597). C. S. niponica var. niponica (M. Tagawa & K. Iwatsuki 3014, P01575937). Spores of Struthiopteris: D. S. castanea (Y. Saito s.n., TNS792168). E. S. fallax (P. Wasowicz & J.M. Gabriel y Galán s.n., MACB 109359); F. S. spicant var. spicant (A. Santos s.n., MACB36152). G. S. spicant var. homophyllum (Barrera s.n., MACB32367); H. S. spicant var. pradae (S. Molino et al s.n., MACB110654). Bar = 30 μm in A–D; 22 μm in E–H.

FIGURE 10. Spicantopsis species in the field. A‒C. Spicantopsis amabilis. A. Habit; B. Detail of the abaxial side of the fertile pinnae, bearing sori; C. Detail of the rhizome scales; D‒E. Spicantopsis hancockii. D. Habit; E. Detail of the abaxial side of the fertile frond, bearing sori; F-G. Spicantopsis niponica var. niponica. F. Habit; G. Detail of the abaxial side of the fertile frond, bearing sori. Pictures: A. Kimitsu, Chiba, Japan, by Keita Watanabe; B & C. Hichiso, Gifu, Japan, Yukihiro Emoto; D & E. Yilan, Taiwan, by Gu Jianye; F. Fujinomiya, Shizuoka, Japan by Harum Koh; G. Izumi, Osaka, Japan, by Tatsuya Fujiwara.

FIGURE 13. Illustration of Spicantopsis niponica. A. Habit of S. niponica var. niponica (J. Umemura 37, P01406521); B. Rhizome scale of S. niponica var. minima (M. Tagawa 853, P00748579, isotype); C. Habit of S. niponica var. minima (M. Tagawa 853, P00748579, isotype); D. Sporangium of S. niponica var. niponica (B. Estébanez s.n., MACB110657); E. Rhizome scale of S. niponica var. niponica (Maximowicz s.n., S2145). Scale: 4 cm in A; 500 µm in B; 4 cm in C; 100 µm in D; 1.5 mm in E.