Spicantopsis amabilis (Makino) 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 : 10

Spicantopsis amabilis (Makino) Nakai ( Figure 9A ; Figure 10A‒C ; Figure 11 ).

Type:— JAPAN. Awa Prov. , Mt. Kiyosumi, 6 April 1896, T. Makino s.n. ( MAK113467 ! photo, lectotype, designated here; MAK115262 ! photo, isolectotype) .; JAPAN. Musashi Prov. , Chichibu, Mt.Yokami, 16 July 1888, T. Makino s.n. ( MAK113468 ! photo, MAK115263 ! photo, syntypes) ; JAPAN. Sabami Prov. , Hakone-tonosawa, 31 December 1893, M. Matsuda s.n. ( MAK113469 ! photo, syntypes) .

Synonyms:— ≡ Blechnum amabile Makino. Spicanta amabilis (Makino) Nakai (1928: 12) . Struthiopteris amabilis (Makino) Ching (1940: 243) . Lomaria amabilis (Makino) Ching (1940: 5) .

Invalid designations: Lomaria crenulata Moore in Hooker & Baker (1867: 180) nom. nud.; Blechnum crenulatum Salomon (1883: 115) .

Plants terrestrial; rhizomes short to more or less long creeping, covered with concolorous, light brown, ovate-lanceolate, membranaceous scales, 1–2 (4) x 2–7 (11) mm, apices acute, margins entire, slightly cucullate; fronds dimorphic; sterile fronds (10) 21 (35) cm long, stipes brown or green (0.5) 2 (10) cm long, laminae linear-lanceolate, coriaceous, (1.5) 3 (5) x 10–17 (30) cm, 1-pinnate or pinnatisect, apex conform, glabrous, rachis brown or green, without adaxial groove, glabrous, pinnae tapering towards base, the middle ones 0.2–0.5 x 0.8–2.5 cm, costae brown or green, glabrous; veins simple or 1-furcate, free, catadromous, ending in submarginal patent hydathodes; fertile fronds about the same length as sterile ones, sometimes shorter, stipes short, green, (1) 3–13 cm, laminae lanceolate, narrower than the sterile ones (1.5–3 cm wide), 1-pinnate, apex conform, rachises brown or green, glabrous, without adaxial groove, pinnae contracted, the middle ones 0.15–0.5 x 0.6–2 cm; sori linear, continuous, forming coenosori on both sides of costa, covering entire length of pinna, not decurrent to rachis; indusia linear, continuous, open to costa, long and coiled; sporangia with 20‒26 arc cells; spores monolete, 30–50 x 50–70 µm, brown, with verrucose-granular perisporium.

Habitat and distribution:— Endemic to Japan, in Honshu, Shikoku (Kochi prefecture) and Kyushu ( Figure 6 ). Grows on rock outcrops in the undergrowth of the mountainous zone, and on rocky slopes and cliffs ( Ohwi 1953, Iwatsuki 1992, Iwatsuki et al. 1995). 300–1700 m.

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

Etymology:— Makino applied the Latin epithet amabile (amiable, pleasant) to this taxon apparently due to its slender and graceful habit. The vernacular, osashida, means “loom-comb fern”, probably because of its resemblance of its fronds to this object.

Vernacular names:— Osashida (オサシダ, Japanese) ( Iwatsuki 1992).

Taxonomic notes:— It differs from the other species of the genus in its rhizome scales, which are membranaceous, concolorous and ovate-lanceolate, whereas S. hancockii and S. niponica show papyraceous, linear, discoloured, sclerotized scales. Spores have a verrucous-granular perispore in S. amabilis , which is irregularly reticulate in the other species.

In the protologue, Makino cites several syntypes ( Makino 1897). We have selected T. Makino s.n. (MAK113467) as lectotype (since it is the one in which the long creeping rhizome, the character Makino considered most distinctive, is most evident) and T. Makino s.n. (MAK115262) as isolectotype.

During our revision we found an undated and apparently old specimen allegedly collected in China (Sichuan) that lies within the morphological circumscription of S. amabilis (E. Faber 1028, NY!). However, we prefer to consider this presence doubtful given the scarce information available (there is no information about county, province, or city) until it can be verified with more precise records.

Aditional specimens examined:— JAPAN. Honshu: Pref . Aomori: Aomori-shi , 18 October 1981, M. Neichi s.n. ( TNS517591 ! photo) ; Imabetsu-machi , 23 October 1977, N. Saitoh ( TNS517423 ! photo) ; Noheji-machi , 18 August 1972, T. Sudo s.n. ( TNS517425 ! photo) ; ibidem, 31 August 31, T. Sudo s.n. ( TNS517674 ! photo) ; Oirase , 4 August 1977, T. Naito s.n. ( MO2697240 !) ; Soma-mura, 28 July 1979, M. Saito & A. Takahashi s.n. ( TNS517424 ! photo) ; Towadako-machi, 31 August 1975, T. Sudo & M. Sudo s.n. ( TNS463340 ! photo) ; ibidem, 13 October 1981, M. Neichi s.n. ( TNS517426 ! photo) ; Pref. Chiba: Kazusa-machi , 1 December 1963, F. Konta 3972 ( TNS924822 ! photo) ; Mitsuishiyama, Kameyama, Kazusa-cho , Kimitsu-gun , 2 July 1966, G. Murata & K. Iwatsuki 580 ( US2424195 !) ; Mt. Nokogiri , 21 March 1924, Y. Yamamoto s.n. ( TAIF125401 ! photo) ; Pref. Fukushima: Yama-gun, Kitashiobaramura, Nakatsugawa Gorge , 9 July 1999, M. Maki et al 997975 ( MO5333264 !) ; Pref. Gifu: Takayama, 1915, J. Faurié 51 ( NY04179844 !) ; Pref. Gunma: Kusatsu, 1877, M. Dickins s.n. ( P01406525 !) ; Hyogo, Kobe city, Mt. Rokko , 20 July 1932, K. Uno 9171 ( TAIF461138 ! photo) ; Pref. Iwate: Kayusa, Mt. Mitsuishi , 13 December 1953, T. Namegata 3819 ( US2138756 !) ; Pref. Kanagawa: Kamkura, 1915, T. Makino 32949 ( US2421838 !) ; Pref. Kyoto: Oohuse, north of Kyoto , 23 September 1951, M. Tagawa 4240 ( US2258980 !) ; Tadasu-no-mori, 10 July 1888, J. Faurié 614 ( P01567610 !) ; Pref. Mie: Miyama. cho, 25 June 1983, J. Haginiwa ( TNS984311 ! photo) ; Utobi, 23 November 1956, K. Seto 6672 ( P01608624 !; C0364698 F!; US2420040 !) ; Pref. Nagano: Kawakami-mura , 6 August 1970, M. Kato 39 ( TNS924821 ! photo) ; Shinano, no date, A. Noguchi s.n. ( MO3155632 !) ; Togakushi , 20 August 1906, M. Weigel s.n. ( P01571952 !) ; Ibidem ( P01406644 !) ; Tokiwa, Omachi-shi, Tokiwa , 26 June 1982, T. Yoshizawa 71566 ( TAIF052352 ! photo) ; Pref. Nara: en route from Mt. Wasamata to Mt. Daifugen, Kamikitayama-mura , Yoshino-gun , 21 September 1986, S. Tsugaru et al 7438 ( MO4012486 !) ; Mt. Shofuken-dake in the Oomine Mountains , 4 August 1962, M. Tagawa 537 ( US2423751 !) ; Shôfuken-dake, 4 August 1962, M. Tawaga & K. Iwatsuki 5129 ( P01575933 !) ; Pref. Niigata: Echigo, Takidani , 24 August 1903, M. Weigel s.n. ( P01406649 !) ; Echigo, Tsugawa , 16 September 1957, M. Togashi 1576 ( MO1734177 !; NY04179843 !; P01406637 !; S1952 !; US2259174 !) ; Itoigawa , 3 August 2009, S. Matsumoto s.n. ( TNS9544305 !) ; Kitauonuma-gun, Irihirose-mura, Ooshirakawa , 22 August 1964, H. Kanai et al. 8966 ( TI00274373 ! photo) ; Shibata-shi , 1 July 1962, F. Konta 402 ( TNS924818 ! photo) ; Ibidem, F. Konta 981 ( TNS924824 ! photo) ; Ibidem , 27 October 1963, F. Konta 3394 ( TNS924823 ! photo) ; Ibidem , 27 August 1966, F. Konta 5698 ( TNS924809 ! photo) ; Ibidem , 1 November 1967, F. Konta 6741 ( TNS924812 ! photo) ; Pref. Saitama: Chichibu-gun, Ohtaki-mura , 9 July 2002, T. Iwata s.n. ( TNS1147001 !) ; Ibidem ( TNS01147002 !) ; Musasi, Ryôgami , 16 September 1957, M. Tagawa 341 ( P01406650 !) ; Pref. Shiga: Umenoki to Kawai, Katsuragawa-mura , Shiga-gun , 22 August 1962, M. Tagawa 8687 ( US2551253 !) ; Pref. Shizuoka: Amagiyugashima-cho, 23 July 1966, F. Konta & M. Wakabayashi 65 ( TNS924811 ! photo; TNS924808 ! photo) ; Fuji-shi , 15 September 1983, T. Sato 4368 ( TNS924830 ! photo) ; Fujinomiya-shi , 16 September 1984 ( TNS924816 ! photo) ; Hamamatsu-shi , 8 August 2000, T. Nakamura s.n. ( TNS1019335 !) ; Misakubo-cho , 10 October 1986, F. Konta 15891 ( TNS924829 ! photo) ; Nyuhjima, Umegashima, Sugura , 9 August 1979, Y. Saiki 1049 ( C2103458 F!) ; Shizuoka-shi , 8 August 1973, F. Konta 10064 ( TNS924810 ! photo) ; Umegashima-mura , July 1964, S. Choshi s.n. ( TNS924813 ! photo) ; Pref. Tochigi, 13 September 1958, M. Tagawa & K. Iwatsuki 1913 ( P01608623 !) ; ibidem ( US2358631 !) ; Pref. Tomaya, Tateyama , August 1913, J. Faurié 51 ( S1953 !) ; ibidem ( P01406643 !) ; ibidem ( P01406645 !) ; ibidem ( P01406646 !) ; Pref. Wakayama: Oto-mura, 11 August 1965, G. Murata & F. Konta 122 ( TNS924814 ! photo) ; Kii, Owase-machi , Kita-muro-gun, 9 October 1956, M. Furuse s.n. ( S1951 !) ; Kii, Yukawa , 25 July 1883, J. Matsumura s.n. ( TI00274283 ! photo) ; Pref. Yamagata: en route from Higashiootori-dam to Awataki-dam, Asahi-mura , Higashitagwa-gun , 18 July 1990, S. Tsugaru 13489 ( MO4296957 !) ; Pref. Yamanashi: Fusiyama, 1864, Tschonoski s.n. ( P01406532 !) . Kyushu: Pref. Kagoshima: Ibusuki-shi , 28 August 1983, T. Yamanaka s.n. ( TNS779978 ! photo) ; Okuchi-shi , 12 January 1969, T. Yamanaka s.n. ( TNS734286 ! photo) ; Satsuma , July 1888, J. Faurié s.n. ( P01567609 !) ; Yakushima Island, Yaku-cho, 12 October 2006, A. Ebihara & M. Ito TF297 ( TNS763339 !). Shikoku. Pref . Kouchi: Tosa, Mt. Kaminomine , 13 April 1930, H. Itô s.n. ( TI00274351 ! photo). Unknown location: no date, no collector ( P01406642 !) ; 20 August 1910, no collector ( US1095353 !) ; 9 February 1910, Christ 94 ( P01406586 !) ; August 1913, J. Faurié s.n. ( P01575934 !) ; 28 August 1898, J. Faurié 1566 ( P01406590 !) ; July 1904, J. Faurié 5612 ( P01406647 !) ; ibidem ( P01406648 !) .

Figures

FIGURE 6. Distribution of Spicantopsis amabilis extracted from GBIF.

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 11. Illustration of Spicantopsis amabilis. A.Habit (Nakamura s.n., TNS01019335); B. Sporangium (J.Faurié 5612, P01406648); C. Rhizome scale (J. Faurié s.n., S1953). Scale: 4 cm in A; 100 µm in B; 1 mm in C.