Spicantopsis hancockii (Hance) Masam., 1936

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 : 12-14

Spicantopsis hancockii (Hance) Masam. ( Figure 9B ; Figure 10 D‒E ; Figure 12 ).

Type:— TAIWAN. In rupibus apertis montis olim ignivomi nunc exstincti Huangkong, juxta Tam-sui, insulae Formosae, alt. 3300 ped. rarissimum, nec alibi, coll. 27 November 1881, W. Hancock 87 (Herb. propr. n. 22183, BM 001045995! photo, holotype).

Synonyms:— ≡ Blechnum hancockii Hance ; Lomaria hancockii (Hance) Baker (1891: 220) ; Spicantopsis niponica (Kunze) Nakai var. hancockii (Hance) Nakai (1933: 183) ; Struthiopteris hancockii (Hance) Tagawa (1952: 192) .

= Lomaria apodophylla Baker (1885: 104) . Type :— NORTH FORMOSA [ TAIWAN]. no date, W. Hancock 87 (US00065538!, holotype; LE01009883 ! photo, isotype).

Plants terrestrial, sometimes saxicolous; rhizomes erect short, covered with linear-lanceolate, discoloured scales, with the centre broadly sclerotized from base to apex, chestnut or dark red, margins light brown, 0.5–1 x (5) 10 (16) mm; fronds dimorphic; sterile fronds (11) 27 (45) cm long, shortly petiolate, stipes light brown or green, 0.5–1 (4) cm long, laminae linear to elliptic-lanceolate, coriaceous, 3‒6 (10) x 10‒30 (45) cm, 1-pinnate, pinnatifid or pinnatisect, apex subconform, abaxial side with small, concolorous, light brown, triangular-lanceolate scales, rachises light brown or green, grooved adaxially, abaxially with scales similar to those on the lamina, pinnae tapering towards the base, the middle ones 0.3‒0.6 x 1.5‒4 (8) cm, tapering towards base, veins free, simple or 1-furcate, catadromous, ending in submarginal patent hydathodes; fertile fronds shorter or about the same size as sterile ones, stipes brownish or green, 1‒3 (7) cm, laminae coriaceous, 2‒5 (8) x 10‒30 (48) cm, 1-pinnate, with scales like those on the sterile lamina and rachis, pinnae more contracted, the middle ones 0.2‒0.4 (0.6) x 1.5‒3 (5) cm, tapering toward base; 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 20‒29 arc cells; spores monolete, 25‒40 x 40‒60 µm, perisporium with granular reticulate surface.

Habitat and distribution:— Taiwan ( Figure 8 ). Relatively dry or moist areas, usually on rocky shaded slopes, terrestrial, sometimes saxicolous. 800‒2900 m ( Iwatsuki 1992, Chiou et al. 1994, Iwatsuki et al. 1995, Kuo 2001, Ebihara et al. 2023).

Chromosome number:— 2 n = 62, diploid ( Fabbri 1965, Nakato 1987). There is another chromosome counting on a recent work ( Nakato et al. 2020), but the specimen is from Japan, where it has been proven that the species does not occur ( Ebihara et al. 2023), so it is probably a specimen of S. niponica .

Etymology:— The taxon honors William Hancock (1847‒1914), Irish botanist and collector of the type specimen during his tenure as customs officer in China and Taiwan (Bionomia 2023).

Vernacular names:— Kuān yè jiá náng jué (Ĩ叶ẍ囊″, Chinese); Haku-un-shida (ハクウンシダ, Japanese) ( Iwatsuki 1992, Faguo et al. 2013). The Japanese name, haku-un-shida, means “fern of the white cloud”, we have no certainty of the reason of this name. The Chinese name, Kuān yè jiá náng jué, means “pod fern”, we are not certain why either.

Taxonomic notes:— Spicantopsis hancockii is very similar to S. niponica , and in fact, authors such as Nakai (1933) or DeVol (1975) considered they could be the same species or, at most, varieties of the same species. However, recent studies consider them as two distinct species (Itwasuki et al. 1995, TPG 2019, Molino et al. 2019b, Ebihara et al. 2023). Different distinguishing characters have been proposed, for example Otaga (1941) and Nakaike (1982) noted differences in the fertile fronds (shorter than sterile in S. hancockii and conversely in S. niponica ), pinna margins (recurved in S. hancockii ), pinna scales (with “marginal hairs” in S. hancockii and without them in S. niponica ) and spore size (larger in S. hancockii ). Ebihara (2016), however, did not find the differences in scales, while he observed marginal excretions in both species, an aspect also seen by us. Molino et al. (2020) found the opposite relationship in the spores (those of S. niponica being slightly larger), and noticed that the sporangia were clearly differentiable, with two posterior cells in the case of S. niponica and no posterior cells at all in the case of S. hancockii . Relative length of sporophylls relative to trophophylls (sporophylls shorter than trophophylls in S. hancockii and the reverse in S. niponica ) has proven useful, however it may be confusing in some immature specimens of S. niponica , with shorter sporophylls ( Ebihara et al. 2023). In these cases, the examination of the sporangium seems the only reliable character ( Molino et al. 2020, Ebihara et al. 2023). 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). In addition, it has been shown that Spicantopsis hancockii does not coincide in distribution with S. amabilis and S. niponica , the former growing only in Taiwan and the other two only in Japan ( Ebihara et al. 2023). It can be confused with Cleistoblechnum eburneum (Christ) Gasper & Salino , with which it coincides in distribution. They can be differentiated because C. eburneum has fronds with very slight dimorphism, the margins of the pinnae are strongly revolute and the laminae are coriaceous, whereas in S. hancockii the fronds are strongly dimorphic, the margins of the pinnae are straight, and the laminae are less leathery than in C. eburneum .

Several authors have considered that S. hancockii is present in the Tokara Islands, Japan, based on different collections ( Hatsushima 1962, Hatusima 1986, Sako 1978, Shiuchi & Hotta 2015, Suzuki et al. 2022). On the other hand, other authors considered that these collections are indeed S. niponica ( DeVol 1975, Kurata & Nakaike 1987, Chiou et al. 1994, Kuo 1997, 2001, Ebihara 2016). Recently, Ebihara et al. (2023) have confirmed by molecular techniques and micromorphology that specimens from the Tokara Islands are indeed S. niponica , confirming that S. hancockii is a strict Taiwanese endemic.

Additional specimens examined:— TAIWAN. Chiayi: Alishan , 25 December 1928, Y. Kudo & S. Suzuki s.n. ( NL996085 ! photo) ; Ibidem , 2‒4 April 1973, C. M. Kuo 3099 ( TAI161616 ! photo; TAI271480 ! photo) ; Ibidem , 2‒4 April 1973, C. M. Kuo 3121 ( TAI161754 ! photo) ; Ibidem , 2 January 1994, S.- J. Moore 16657 ( TAIF486372 ! photo; TAIF486373 ! photo) ; Ibidem , 1 March 2008, P.- F. Lu 15930 ( TAIF303877 ! photo) ; Ibidem , 2 July 2014, Tashan et al. s.n. ( TNS1219667 !) ; Alishan, Arisan , 22 July 1970, T. Iwasaki s.n. ( TNS832514 !) ; Alishan, Mienyueh, T.- C. Hsu 9408, 13 July 2017 ( TAIF509453 ! photo) ; Alishan, Mt. Ta , 1 March 2008, C.- W. Chen Wade 564 ( TAIF389749 ! photo) ; Alishan, Zhaoping, Mt. Ta , 2 July 2014, Y.- H. Chang 20140702-011 ( TAIF453698 ! photo) ; Mt. A-li, 25 October 1962, M. T. Kao & C. S. Feung 153 ( TAI004260 ! photo) . Hsinchu: Jianshih, Meiyuan, Jianshih , Mt. Niaotsui , 12 July 2009, P.- F. Lu 18523 ( TAIF335193 ! photo) ; Jianshih, Mt. Litung , 6 September 2009, C.- W. Chen Wade 969 ( TAIF397475 ! photo) ; Kuan-wu, 99 Hostel to Ta-pa-chien Shan , 4 April 1972, C. C. Hsu & C. S. Kuoh 13980 ( TAI147992 ! photo) ; Ta-pa-chien Shan , 10 August 1972, C. M. Kuo 1756 ( TAI153035 ! photo; TAI159196 ! photo) ; ibidem, 11 August 1972, C. S. Kuoh 3321 A ( TAI157143 ! photo) . Hualien: Erhtzushan , 6 September 1983, T.- T. Lin & Y.- F. Chen 7153 ( TAIF216160 ! photo) ; Jeisui logging trail, 16 November 2007, P.- F. Lu 14917 ( TAIF291133 ! photo) ; Mt. Muh-kwa , 24 July 1961, M. T. Kao 4140 ( MO3692478 !; TAI004256 ! photo; TAI126342 ! photo; TAI157281 ! photo) ; Muh-Kwa , 24 July 1961, M. T. Kao 4141 ( TAI004259 ! photo) ; Taipingchichi , 20 September 1999, M.- L. Weng 1598 ( TAI150756 ! photo; TAI261730 ! photo) . Illan: Datong, Provincial Rd. No. 7 A, 27 September 2007, F.- W. Li FWL560 ( TAIF371276 ! photo) ; Datong, Ssuyuanyaku , 18 October 1994, Y.- H. Chang 60 ( TAI278160 ! photo) ; ibidem, 11 August 2012, Y.- S. Chao 1965 ( TAIF435569 ! photo) ; Datong, Taipimgshan ( Tsubao ), 28 March , 1930, S. Suzuki 3914 ( TAI119360 ! photo) ; Mt. Taiping , 20 March 1963, C. S. Feung & M. T. Kao 366 ( US 2635362!; TAI004261 ! photo) ; ibidem, 4 March 1966, C. C. Chuang & M. T. Kao 4136 ( TAI004258 ! photo) ; ibidem, 4 March 1967, M. T. Kao 6794 ( TAI004257 ! photo) ; Nan’ao, Mt. Nanhu , 27 September 2007, E. Schuettpelz et al. 1175 B ( TAIF288823 ! photo) ; Nan-ao, Mt. Tung , 4 November 2012, P.- F. Lu 24831 ( TAIF403619 ! photo) ; Nan’ao, Taipingshan , 7 August 1981, W.- L. Chiou s.n. ( TAIF166973 ! photo) ; Ibidem , 28 January 1994, J. C. Wang et al. 9308 ( TAIF79814 ! photo) ; Ibidem , 3 September 2016, R. Knapp 4311 ( P02435597 !) ; Ibidem , 5 December 2005, R. Knapp & J.- Y. Huang 239 ( P02436496 !) ; Nan’ao, Taipingshan, National Forest Recreation Area, Cueifong Lake Circle Trail , 20 June 2008, A. Ebihara et al. s.n. ( TNS776516 !) ; Nao’ao, Tsuifeng lake , 28 February 2004, P.- F. Lu 7611 ( TAIF202731 ! photo) ; Nan-hu-ta-shan, 16 October 1977, J.- C. Wang 1723 ( TAI198587 ! photo) ; Nan-shan , April 1944, unknown collector 417 ( TAI161743 ! photo; TAI161747 ! photo) ; Taipingshan, 14 May 1992, S. F. Huang 4771 ( TAI222807 ! photo). Kaohsiung: Chienchin-Kuaishan, Miharasi- Hinokiyama , 9 December 1937, Yamamoto 615 ( TAI004265 ! photo) ; Shih-shan , 23 January 1972, C. M. Kuo 664 ( U0043511 ! photo; TAI161755 ! photo) ; ibidem, 14 November 1972, C. M. Kuo 2456 ( TAI162439 ! photo) ; ibidem, C. M. Kuo 6507 ( TAI146786 ! photo). Miaoli: Taian, Leshan logging trail, 27 May 2004, P.- F. Lu 7940 ( TAIF210083 ! photo). Nantou: Dayuling , 14 July 2007, R. Knapp 1413 ( P02437438 !) ; Luantashan , 28 April 1922, S. Kudo s.n. ( TAIF126083 ! photo) ; Sinyi, Chunta logging trail, 27 November 2012, Y.- H. Chang 20121127-003 ( TAIF444888 ! photo; TAIF444889 ! photo) ; Ibidem , 9 September 2013, H.- C. Hung s.n. ( TAIF452653 ! photo) ; Sinyi, Kuankao, Tuikuan , 15 December 2011, T.- C. Hsu 5195 ( TAIF387835 ! photo) ; Sinyi, Shuanglung logging trail, 12 September 2012, H.- C. Hung & Y.- H. Chang 20120912-004 ( TAIF408677 ! photo). Pingtung: Gaoshu, Dr. Cecilia Koo Botanic Conservation Center, 5 February 2014, L.- Y. Kuo & C.- M. Chen 3927 ( TAIF447324 ! photo) ; Ibidem , L.- Y. Kuo & C.- M. Chen 3928 ( TAIF447323 ! photo) ; Peitawushan , 7 February 1987, B.-J- Wang 12942 ( TAIF162091 ! photo; TAIF162092 ! photo) ; Peitzasuahzn , 27–29 January 1978, Y. C. Jeng 2744 ( TAI277663 ! photo) ; Taiwu, Kuaiku, Tawu Shrine , 8 October 2006, P.- F. Lu 12661 ( TAIF184495 ! photo) ; Wutai, Hsiaokuei lake, 18 February 2015, P.- F. Lu 27885 ( TAIF460900 ! photo). Taichung: Baxian Mountain [Eight Immortals Mountain], 24 October 1929, S. Suzuki 2960. ( TAI067582 ! photo; TAI161746 ! photo) ; Heping , 710 logging trail, 27 September 2007, Y.- H. Chang ( TAIF296602 ! photo) ; Heping, Mt. Loyehwei , 25 April 2015, P.- F. Lu 28041 ( TAIF465472 ! photo) ; Ibidem , 1 August 2015, P.- F. Lu 28511 ( TAIF467412 ! photo) ; Heping, Ssuyuanyakou, Mt. Nanhu , 1 July 2009, Y.- H. Chang 20090701-021 ( TAIF400098 ! photo) ; Nanhu Dashan , October 1928, S. Sasaki s.n. ( TAI004270 ! photo) ; Mt. Hassenzari , 23 October 1929, S. Suzuki s.n. ( TAI007530 ! photo) ; Pa-Shen-Tun to Mt. sin Shan , 7 August 1959, T. C. Huang 1321 ( TAI004267 ! photo) ; Pa-shien-shan, 5 November 1969, T. C. Huang 1336 ( TAI12776 ! photo) ; Pahsinshan , 7 August 1959, Huang 1321 ( TAI004272 ! photo) ; Pahsinshan Logging Station , 7 August 1959, T. I. Chuang et al. 2751 ( TAI004262 ! photo; TAI004264 ! photo) ; Ssuyuan, 11 January 2001, J.- H. Li & C.- K. Liou 399 ( TAIF153000 ! photo; TAIF153001 ! photo). Taipei: Beitou , 18 January 2014, R. Knapp 3338 ( P02439073 !) ; Beitou, Chihsing , 28 February 2003, P.- F. Lu 5529 ( TAIF187192 ! photo) ; Ibidem , 19 September 2009, Y.- H. Chang 20090919-004 ( TAIF366933 ! photo) ; Ibidem , 1 June 2013, P.- F. Lu 25597 ( TAIF418892 ! photo) ; Chihsingshan , 16 July 1974, C. E. DeVol 1202 ( TAI187939 ! photo) ; ibidem, 14 June 2000, P.- H. Lee 300 ( TAIF115743 ! photo) ; Ibidem , 12 January 2002, P.- F. Lu 2908 ( TAIF155311 ! photo; TAIF155312 ! photo) ; Mt. Seven Star , 11 October 1963, M. T. Kao 5466 ( TAI004266 ! photo) ; Sitisei, Chihsingshan , 2 September 1926, S. Suzuki s.n. ( TAI004269 ! photo) ; Peichatenshan , 28 September 1984, R. T. Li 3583 ( TAI196166 ! photo) ; ibidem, R. T. Li 3584 ( TAI196165 ! photo) ; ibidem, R. T. Li 3585 ( TAI196164 ! photo) ; Taiping , 20 February 1963, Feung & Kao 336 ( TAI004261 ! photo) ; Tsi-hsin-shan, 15 November 1972, C. P. Ling 2468 ( TAI148816 ! photo) ; Tsi-hsin-shan (Suen Star Mt.) 28 January 1924, no collector s.n. ( TAI132647 ! photo). Taitung: Beinan, Chihpen logging trail, 6 December 2012, C.- F. Chen 4361 ( TAIF423999 ! photo) ; Taiwu , 2 June 2014, R. Knapp 3430 ( P02439164 !; P02439165 !) ; Yanping, Yenping logging trail, 10 December 2002, P.- H. Lee 2326 ( TAIF182691 ! photo; TAIF182691 ! photo). Taoyuan: Fushing, en route from Takuanshan Forest Reserve to Tamanshan, 9 April 2002, W.- C. Leong 2786 ( MO6428317 !) ; Fusing, Howi Giants Group, New Taipei City , Sansia , Mt. Peichantien , 7 August 2011, C.- W. Chen Wade 1658 ( TAIF397367 ! photo) ; Fushing, Mt. Fufu , 19 June 2010, T.- C. Hsu 2838 ( TAIF392212 ! photo) ; Fushing, Mt. Peichatien , 1 November 2014, P.- F. Lu 27372 ( TAIF456353 ! photo) ; Kaohsiung city, Chuyunshan logging trail, 5 February 1992, B.- J. Wang 18990 ( TAIF165847 ! photo; TAIF165848 ! photo; TAIF165849 ! photo; TAIF165850 ! photo; TAIF165851 ! photo). Unknown location: 1 August 1906, H. Shakurai s.n. ( P01406577 !) .

Figures

FIGURE 8. Distribution of Spicantopsis hancockii 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 12. Illustration of Spicantopsis hancockii. A. Habit (Y.-H. Chang 60, TAI278160); B. Sporangium (R. Knapp 3338, P02439073); C. Rhizome scale (R. Knapp 3430, P02439164). Scale: 4 cm in A; 100 µm in B; 500 µm in C.