广布铁角蕨 guang bu tie jiao jue

Asplenium leiboense Ching; A. sarelii Hooker var. anogrammoides (Christ) Tagawa; A. sepulchrale Hooker, p.p.

Plants 10-30 cm tall. Rhizome short, erect or ascending, apex scaly; scales dark brown to black, narrowly triangular, 3-8 × 0.3-0.7 mm, margins denticulate; scale base hyaline, cordate, with numerous yellow-brown, unicellular hairs 1-5 × ca. 0.02 mm, similar to root hairs. Fronds caespitose; stipe 7-15 cm, semiterete, dark brown abaxially or becoming green toward rachis, upward green, adaxially green, shallowly sulcate with supravascular ridge, with similar scales as on rhizome, reduced hairlike scales toward rachis or subglabrous; lamina ovate, 6-13 × 2.5-7(-8) cm, base truncate, apex acute, tripinnatifid-tripinnate; pinnae 8-12 pairs, basal pairs subopposite to alternate, stalk 2-4 mm, sulcate adaxially; several basal pinnae pairs ± equal in length, ovate-triangular, 1.5-3 × 0.5-1.5 cm, base nearly symmetrical, truncate to broadly cuneate, 2-pinnatifid-bipinnate, apex acute; pinnules 4-6 pairs, anadromous, acroscopic and basiscopic pinnules ± equal in size, narrowly triangular to ovate, 5-10 × 3-6 mm, base asymmetrical, acroscopic side truncate, basiscopic side cuneate, largest pinnules stalked, others decurrent on costa, pinnatipartite-pinnate, apex acute; segments 2 or 3 pairs, linear to narrow elliptic, apex with 2 or 3 teeth. Costa sulcate adaxially, without median ridge, veins slightly raised adaxially, anadromously forked. Fronds firmly herbaceous, green to grayish green, lamina subglabrous, average stomatal guard cell length 48-52 µm; rachis green, semiterete, abaxially green or base castaneous becoming green toward apex, adaxially sulcate with raised median supravascular ridge, with reduced scales or subglabrous. Sori 2-4 per ultimate segment, median to subterminal on acroscopic vein, subconfluent at maturity, subelliptic, 1-2.5 mm; indusia grayish green, subelliptic, membranous, entire, opening toward costule or central veinlet, persistent. Spores brown, perispore lophate (cristate), average exospore length 33-36 µm. Plants sexual, allotetraploids: 2n = 144.

Wet cliffs, in crevices of limestone rocks, on buildings and walls; 300-2800 m. Anhui, Fujian, Guangdong, Guizhou, Hebei, Hubei, Hunan, Jiangsu, Jiangxi, Jilin, Liaoning, Ningxia, Shaanxi, Shandong, Shanxi, Sichuan, Yunnan, Zhejiang [?India, Japan, Korea, Vietnam].

Asplenium anogrammoides is an (allo)tetraploid (Kurita, J. Jap. Bot. 35: 269-272. 1960; Mitui, Sci. Rep. Tokyo Kyoiku Daigaku, B, 13: 285-333. 1968) and originated via chromosome doubling in the sterile hybrid between A. sarelii and A. tenuicaule (Lin & Sleep in K. H. Shing & K. U. Kramer, Proc. Int. Symp. Syst. Pterid. 111-127. 1989; Murakami et al., Amer. Fern J. 89: 232-243. 1999; Viane & Reichstein, Pterid. New Millennium, 73-105. 2003; Wang, Acta Bot. Sin. 45: 1-14. 2003). Their morphological similarity led to much confusion and misapplication of the name A. sarelii to this taxon (also called "tetraploid sarelii") (e.g., Christensen, Acta Horti Gothob. 1: 41-110. 1924; Ogata, Icon. Fil. Jap. 1: t. 8. 1928; Tardieu, Asplén. Tonkin, 46. 1932; Ching, Icon. Fil. Sin. 3: t. 111. 1935, p.p.; Fl. Pl. Herb. Chin. Bor.-Or. 1: 37. 1958; De Vol, Mus. Heude. Notes Bot. Chin. 7: 93. 1945, p.p.; Tagawa, Col. Ill. Jap. Pterid. 151. 1959; Kurata, J. Geobot. 11: 66-69. 1962; Mitui, loc. cit. 1968; Bull. Nippon Dental Coll., Gen. Educ. 4: 221-271. 1975; Kurata & Nakaike, Ill. Pterid. Jap. 2: 164-173. 1981; Ding & Gao, Fl. Henan 1: 67. 1981; Sleep & Reichstein, Candollea 39: 675-691. 1984; Ching & S. H. Wu, Acta Phytotax. Sin. 23: 1-10. 1985; Jiang, Fl. Anhui 1: 136. 1985, p.p.; Bir, Biol. Indian Pterid. 215-221. 1987; Jamir & Rao, Ferns Nagaland, 289-290. 1988; H. S. Kung, Fl. Sichuan. 6: 377. 1989; Chen, Fl. Shandong 1: 96. 1990; Li, Fl. Liaoning 1: 77. 1990, p.p.; L. K. Lin, Fl. Fujian. 1: 131. 1991; Iwatsuki, Ferns Jap. 146. 1992; Fl. Jap. 1: 104. 1995; Murakami et al., loc. cit.; S. H. Wu, FRPS 4(2): 98. 1999, p.p.; P. S. Wang & X. Y. Wang, Pterid. Fl. Guizhou, 138. 2001, p.p.; Li et al., Fl. Hunan 1: 287. 2004, p.p.; G. F. Zhang, Fl. Yunnan. 20: 669. 2006). Moreover, less-divided plants of A. anogrammoides are similar to large plants of A. pekinense (the autotetraploid arisen from A. sarelii).

On the other hand, the name Asplenium anogrammoides has been misapplied (e.g., Komarov, Izv. Imp. S.-Peterburgsk. Bot. Sada 16: 145-151. 1916; Komarov & Klobukova-Alisova, Opred. Rast. Dal’nevost. Kraia 1: 82. 1931; Fomin, Fl. Sibir. Orient. Extremi 5: 152-154. 1930; in Komarov & Iljin, Fl. URSS 1: 68. 1934; Ching, loc. cit.: 37. 1958) to S Siberian and Mongolian A. tenuicaule var. subvarians.

Yellow-brown (root)hairs are present on the rhizome, stipe base, and scale base of Asplenium altajense, A. anogrammoides, A. pekinense, A. sarelii, A. tenuicaule, and other related taxa, and thus cannot be used to distinguish them. The average exospore and stomata length are the most reliable characters discriminating between diploid and tetraploids in this complex. Asplenium anogrammoides can often be separated from A. pekinense by its largely not-reduced lamina base and its ovate-triangular (vs. deltoid) pinnae with stalks 2-4 mm (vs. 0.5-2 mm).

Where Asplenium anogrammoides grows together with A. tenuicaule, their sterile triploid hybrid, A. ×mitsutae, is common. Asplenium ×mitsutae Viane & Reichstein, nothosp. nov. Type: China. Sichuan: Qingcheng Shan, ca. 70 km WNW of Chengdu, mixed evergreen forest with Rhododendron, on wall inside temple yard together with A. anogrammoides and A. tenuicaule, 1050 m, 16 Sep 1988, Viane 4046-B (= TR-7177) (holotype, GENT). Planta hybrida, inter parentes A. anogrammoides et A. tenuicaule quoad divisionem laminae atque pinnularum segmentorumque formam necnon dimensiones cellularum accessoriarum stomatum intermedia, ab eis sporis abortivis necnon chromosomatum numero triploideo (2n = 108, meiose bivalentibus univalentibusque 36) differt. Meiotic chromosome behavior in this hybrid shows that A. anogrammoides contains one chromosome set of A. tenuicaule. This hybrid is named after S. Mitsuta, who collected it for us in Japan with several other living plants in 1985. Asplenium sarelii var. magnum H. S. Kung, described from Sichuan, is probably the same hybrid combination.

"Asplenium wudangense" (Z. R. Wang & X. Hou, Acta Bot. Sin. 45: 4. 2003) belongs here but was not validly published because no Latin description or diagnosis, or reference to such, was provided (Melbourne Code, Art. 39.1).