Pteris grevilleana Wall. ex J.Agardh

Chao, Yi-Shan, Liu, Ho-Yih & Chiou, Wen-Liang, 2015, Taxonomic revision of the Pteris cadieri complex (Pteridaceae), Phytotaxa 230 (2), pp. 130-150 : 139-148

3. Pteris grevilleana Wall. ex J.Agardh

3a. Pteris grevilleana Wall. (1831: n. 2680) ex J. Agardh (1839: 23). Type :— INDIA. Silhet, Wallich 2680 (lectotype E-00257673!, Figure 6 , here designated; isolectotype K-H2006101585_1!, B-20-0139511!)

Deciduous or evergreen. Rhizomes short, erect, apex scaly; scales linear lanceolate, 1–2 mm long, 0.2–0.5 mm wide, bicolorous, centers dark brown, margins light brown, entire, apex long-acuminate. Fronds clustered, 10–70 cm long, monomorphic, laminae larger and segment narrower on sterile one, stipe of fertile one about twice length as laminae; stipes dark-red or adaxially green, 5–55 cm long, sparsely scaly at base, grooved on the adaxial side; laminae ovate, 1/4–1/2 length of frond, 3–20 cm long, 4–18 cm wide, regularly bipinnatifid; basal pinnae incurved or straight, with exaggerated basiscopic pinnules, terminal pinna similar to lateral ones, 1–3 pairs lateral pinnae; rachises winged and pinnae decurrent along distal stipes; pinnae lanceolate, pectinate, green throughout, 7–13 cm long, 2–5 cm wide, straight or incurved, sessile or short-petiolate, apex caudate or acute; segments of pinnae oblong, slightly oblique to falcate, 3–8 mm wide, margins almost entire to serrulate; sinuses 1–2 mm wide; sori along pinna margins; spores tetrahedral if normal, globose or ellipsoidal if aborted, tan.

2 n = 58, 87, 116 (diploid, triploid, and tetraploid) ( Chao et al. 2010, 2012a, b).

Ecology and distribution:— Southeastern Bhutan, Bangladesh, Cambodia, China, India, Indonesia, Malaysia, Myanmar, Nepal, Philippines, Taiwan, Thailand, and Vietnam; in shaded wet places, under evergreen broad leaf forests, below 1,000 m in elevation.

Etymology:— Named for Robert Kaye Greville, a British botanist, 1794–1866.

Chinese name:—AEṂẶŖdz.

Additional specimens examined: — CAMBODIA. Cocjnchina. s. d., C. Gaudichaud s.n. (B). CHINA. Fujian: Nanjing, Letu, P. F. Lu 17801 (TAIF). Hainan: C. Wang 35869 (HUH, NY); Diaoluoshan, Baishuiling, Y. S. Chao 1257, 1258 (TAIF); Jianfengling, Y. S. Chao 1755 (TAIF); Po-ting, F. C. How 73320 (HUH); Tai Pin, J. L. Gressitt 1115 (E, HUH, UC); Shiqing, Y. S. Chao 1250–1252 (TAIF); Mt. Wuzhi, Y. S. Chao 1204–1207 (TAIF), Wuzhishan Fern Survey 287 (HUH), 368, 561 (HUH); Yaichow, H. Y. Liang 62677 (NY); Mt. Hung Ho, Tsang & Fung 580 (K), 18114 (HUH, K, NY). Hongkong: A. Herry 323 (P), E. Faber 118323 (P). Kouangsi: Long Tcheou, 4 Mars 1900, M. Tanant s.n. (P). Kwangtung: Renhua, Danxiashan, July 7, 2007, X. Y. Dong s.n. (TAIF); Shenzhen City, Wutongshan, Y. S. Chao 1309, 1310 (TAIF); Fang Cheng, W. T. Tsang 26559 (E, HUH, K). Yunnan: Hekou, Nanxi, Nanxi Farm, Y. S. Chao 1319–1322 (TAIF), Nanxi, Y. S. Chao 1333 (TAIF); Xishuangbanna, Shing, Lin & Zhang 6877 (HUH, MICH). INDIA. Assam: Nassibur Forest, Feb 1891, G. Mann s.n. (B); Upper Dehing Reserve Forest, April 1877, G. Mann s.n. (P), Feb . 1891, G. Mann s.n. (P); Lakhimpur Dist., April 1887, G. Mann s.n. (NY); Cachar, June 1, 1873, R. L. Keenan s.n. (K); Sibsagar Dist., Thausi Forest, Jan. 1888, G. Mann s.n. (E, K, US); Margerta, C. B. Clarke 37900 (E, K, P); Namchung, C. B. Clarke 37950B (E). Eastern Circle: Shilong, G. Panigoahi 18800 (K). Meghalaya: Garo Hill, C. B. Clarke 42995 (E, K). Upper Assam: Dec. 1887, G. Mann s.n. (K). JAPAN. Kyushu, Kagoshima: Yakushima Isl., S. Kurata 6170 (US), Jan. 30, 1966, K. Ikuta s.n. (TI); Kumage-gun, Yaku-cho, K. Iwatsuki & M. Kato 92 (KYO), along Odakumi River, S. Mitsuta 12531 (KYO. MICH), near Kurio, H. Hashimoto 123 (KYO), along Tainoko River, Aug. 7, 1936, N. Takaki s.n. (KYO), along Odakumi-gawa River, T. Tagawa & K. Iwatsuki 3197 (KYO), along Hanaage-gawa River, T. Tagawa & K. Iwatsuki 3161 (KYO). Okinawa: Ishigaki Isl., H. Takamine 337 (KYO); Mt. Omoto-dake, 7 Aug 1933, S. Tawada s.n. (B); Iriomote Isl., Mt. Haimi-dake, T. Tagawa & K. Iwatsuki 4692 (KYO). MALAYSIA. Pahang: Tembeling, R. E. Holttum 24726 (MICH). Selangor: B. Molesworth Allen 1688 (US). Kampong Major: Sungei Nerus, Kuala Terengganu, J. Sinclair & Kiah bin 40911 (HUH), J. Sinclair 8697 (E). Kuala Kangsar: Mt. Bubu, B. Molesworth Allen 4737 (HUH). Sabah, Sarawak, C. J. Brooks 14 (MICH); Ranau Dist., Crocker Range, Bukit Lugas, J. H. Beaman 10587b (MICH), J. & M. S. Clemens 28172 ( US); Mt. Kinabalu, D. L. Topping 1605 (HUH, MICH). MYANMAR. H. Young 59 (MICH). TAIWAN. Hualien County: Yuemeishan, Feb . 5, 2009, C. W. Chen s.n. (TAIF). Ilan County: Y. Yamaonto 1494 (TAI); Mt. Oobi, Suoo-gun, April 28, 1938, G. Masamune s.n. (TAI); Jiaosi Township, Yuehmeikeng Waterfall, P. F. Lu 15202, 1427 (TAIF); Fushan, Y. H. Chang 1816 (TAI), Y. S. Chao 1154 (TAIF); from Taiyang Pond to Leng Pond, Aug. 31, 2008, P. F. Lu s.n. (TAIF). Kaohsiung City: Sanping to Nanfengshan, C. S. Kuoh 4050 (TAI); Sanping, T. Y. Yang 67 (TAI). Keelung City: Faurie 126 (P); Chingjenhu, P. F. Lu 11381 (TAIF), C. C. Liao 1121 (HUH); Haimentienhsien, M. F. Kao 3830 (TAI), Aug. 30, 2008, C. W. Chen s.n. (TAIF); Hsingahan Dam, S. F. Huang & K. C. Yang 3788 (TAI); Tona, Kizan-gun, M. Tagawa 1903 (KYO). Nantou County: Shueili Township, Erhpingtzu, P. H. Lee 3173 (TAIF); Yuchih Township, Lienhuachih, J. C. Lin 183 (TAIF), P. F. Lu 7101 (TAIF), P. H. Lee 493 (TAIF), Y. S. Chao 1038–1041, 1046, 1166–1168 (TAIF). New Taipei City: Oct. 22, 1899, K. Miyake s.n.; Chungho Dist., H. L. Chiang 2694 (TAIF); Aoti, S. F. Huang 2937 (TAI), Haihshan Menghu, C. M. Kuo 15462 (TAI); Pinglin Dist., C. H. Chen et al. 3131 (TAIF), Gupoliao River, Y. S. Chao 1075 (TAIF), Mt. Chiuchiungken, Jan. 3, 2009, P. F. Lu s.n. (TAIF); Rueifang Dist., Chintzyupei, Nov. 15, 1990, S. Y. Lu s.n. (TAIF); Shenkeng Dist., Tukuyueh, P. F. Lu 11702 (TAIF); Shihmen Dist., Chienshanhu Trail, Y. H. Chang 20080620-4 (TAIF); Shuangsi Dist., C. T. Lu 414, 5141, 5149 (TAIF); Sijhih Dist., Do 555 (TAI), Hsinshan-menghu, C. Y. Hu 756 (TAIF), Y. H. Chang 4065 (TAIF), Y. S. Chao 1063, 1064 (TAIF); Sindian Dist., I. Shimozawa 556 (TAI), Bunzangun, April 4, 1933, Do s.n. (KYO), M. Tagawa 239 (MICH), Kannai, Aug. 12, 1930, S. Sasaki s.n. (TAI), Wantan, Nov. 22, 1936, H. Shimizu 3240 (TAI), H. Shimizu 2889 (TAI), Agyoku, Togo-kei, Suman, Suzuki-Tokio 13832, 14777 (TAI), M. Tagawa 199, 239 (KYO), Mt. Shihtou, P. F. Lu 5390, 5462 (TAIF), Inhotong, Y. H. Chang 722 (TAI); Wulai Dist., H. M. Chang 3065, 3066 (TAIF), Wulaishan, Y. S. Chao 768 (TAIF); Taipei-Ilan Highway 49.5–50K, B. J. Wang 15316 (TAIF); Maokung, Nov. 24, 2007, P. F. Lu s.n. (TAIF). Pingtung County: Bankinsing, A. Herry 1495 (K, NY); Hengchun Township, Mt. Laofo, P. F. Lu 7718 (TAIF); Kaoshu, Tachin Waterfall, Y. T. Jeng 269 (TAI); Manchou Township, Lanjen Stream, B. J. Wang 7843, 7873 (TAIF), Y. S. Chao 726, 728 (TAIF), Nanjenshan, C. M. Kuo 1990 ×2 (TAI), P. F. Lu 11416 (TAIF), R. Moran 5447 (E, HUH, TAIF, UC, US), Y. S. Chao 770 (TAIF; Mutan, H. Ohashi et al. 13740 (TAI); Shihzih Township, Shouka, P. F. Lu 11061, 1201, 1202 (TAIF), B. J. Wang 13527 (TAIF), Feb . 7, 2009, C. W. Chen s.n. (TAIF), Bei-jih-shan, Kuo & Yu 14334 (HUH), Shouchia, July 10, 2008, Y. H. Chang s.n. (TAIF), Fengkang, M. J. Jung 2287 (TAIF), Kaoshihchiuchih, Y. S. Chao 707, 708, 710, 721, 1065 (TAIF), Maotzushan, Y. S. Chao 701, 702 (TAIF), Luliao River, Dec. 9, 2007, L. Y. Kuo s.n. (TAIF). Taichung: Rengechi, A. T. Hsieh & M. T. Kao 73 (TAI), Dec. 30, 1925, S. Sasaki s.n. (TAI), March 3, 1927, S. Sasaki s.n. (MICH), Sinkogun, Kusuhara, Suibotei, Suzuki-Tokio 19666 (TAI). Taipei City: Dadaoshan, H. C. Ho 40 (TAI), Hsiangshan, M. F. Kao 866 (TAI); Hu-Shan, M. F. Kao 1785 (TAI); Shihlin Dist., Neishuanghsi, M. L. Weng 314 (TAI), P. F. Lu 10462 (TAIF); Hsiaokenghsi Trail, P. F. Lu 2351 (TAIF); Taluntoushan, Y. H. Chang 3896 (TAIF); Mt. Hsiang, Y. M. Huang 643, 644, 1169 (TAIF); Maokung, Y. S. Chao 769 (TAIF), Muchihshan, C. C. Chang 114 (TAI), April 15, 2000, S. C. Wu s.n. (TAI); Yangmingshan, M. F. Kao 286 (TAI). Taitung County: Chihpen, B. J. Wang 15619 (TAIF); Taimali Township, Chinlung, S. W. Chung 6412 (TAIF); Donghe Township, Taiyuan, B. J. Wang 15134 (TAIF); Fenggang river, Jan. 3, 1938, I. Simozawa s.n. (TAI); Taito-gun, near Aroe, M. Tagawa 2453 (K, KYO, MICH, P), 2455, 2461 (KYO), 2467 (KYO, MICH), near Syussuiha, M. Tagawa 2474 (KYO), near Tyatyagatoan, M. Tagawa 2449 (KYO, MICH); near Tyokakurai, M. Tagawa 2451 (KYO, MICH), 2656 (KYO, MICH), 2660 (KYO); Tawu, H. Shimizu 3795 (TAI). Taoyuan County: Haiduan Township, Tianlong Trail, Y. S. Chao 882 (TAIF); Luye Township, Taipingshan, Y. S. Chao 1364, 1365 (TAIF); Lichia Forest Trail, July 27, 2008, P. F. Lu s.n. (TAIF); Lungtan Township, Mt. Shihmen, S. C. Liu & H. M. Chang 705 (TAIF). THAILAND. E. Smith 2362 (K). Chaiyaphum: Khon Kaen, Phu Khieo Game Reserve, G. Murataet al., 49542 (HUH). Huey Ya: R. E. Holtham 11249 (K). Nakawn Sritamarat: Mt. Khao Luang, T. Smitinand 1002 (K); Mt. Khao Luang National Park, Groong Ching Falls, J. F. Maxwell 86-576 (HUH). Nakhon Si Thammarat, at middle elevation of Khao Luang, K. Iwatsuki et al. 8435 (KYO). Phetchabun: Phu Hin Rongkla National Park, April 17, 2008, P. F. Lu s.n. (TAIF). Prachin Buri: Khao Yai National Park, E. Hennipman 3979 (KYO, B). Saraburi: Khao Yai National Park, T. Shimizu et al. 18195 (KYO). Udawn: Loey, Phu Luang, M. Tagawa et al. 1817 (KYO). VIETNAM. C. I Peng 21853 (TAIF). Bac Giang: M. Colani 1597 (MICH, P, UC), 1925, M. Colani s.n. (HUH, TI). Ouonbi: M. Balansa 56 (P), 54 (HUH). Quang-Bihn: L. -M. Cadière 78 (P). Quang Ninh: Dam-ha, Lung Wan Village, W. T. Tsang 29986 (E, HUH); Ha Coi, Shui Mei village, W. T. Tsang 29293 (E, HUH, K); Tong Fa market, W. T. Tsang 29532 (B, E, HUH, K); Kau Nga Shan and vicinty Tien-yen, W. T. Tsang 30599 (E, HUH). Quang Tri: Dakrong, Trieu Nguyen Commune, Y. S. Chao 1595–1602 (TAIF, VNMN). Thai Nguyen: Jan. 6, 1926, M. Colani s. n. (NY); Aou-Du, Eberhardt 5185 (P); Don- Du, collector unkown, 1189 ( US). Thua Thien-Hue: L. -M. Cadière 84 (P). Tonkin: R. P. Bon 2453 (P); Euvirons de lu Phap, Tans les bespuss, B. Balansa 1978 (B, K, MICH, P). Tourane: C. Gaudichaud 95 (P). Vinh Phuc: Dakrong Nature Reserve, Y. S. Chao 1582–1586 (TAIF, VNMN); Tam Dao, Y. S. Chao 1608, 1609 (TAIF, VNMN), Mai 18, 1908, Eberhardt s. n. (P); Tam Dao Town to Cai Keng Road and Bac Thai Road, Y. S. Chao 1626–1632, 1645–1648 (TAIF, VNMN).

3b. Pteris grevilleana Wall. ex J.Agardh forma ornata (Alderw.) Y.S.Chao, H.Y.Liu & W.L.Chiou , stat. nov.

Pteris grevilleana Wall. ex J.Agardh var. ornata Alderw. (1909: 364) . Type:—Borneo, Cult. in Hort.Bog. 1907, Alderwerelt van Rosenburgh s. n. (lectotype BO-1526493!, designated by Chao et al. 2012a).

Distinguished from typical variety by pinnae with whitish or pale, grayish-green central bands on adaxial surface.

2 n = 58, 116 ( Chao et al. 2012a).

Ecology and distribution:— China (Guangdong and Guangxi), Malaysia, India, Indonesia, Myanmar, the Philippines, Taiwan, Thailand, and Vietnam; in shaded places, under evergreen broadleaf forests, below 1,000 m in elevation.

Chinese name:—ŮḢAEṂẶŖdz.

Notes:— Three sheets of syntype specimens of Pteris grevilleana were deposited at B, E, and K, respectively. We here designate the specimen at E as the lectotype because it is the only one with rhizomes and, thus, provides the most complete information.

The Chinese common name for Pteris grevilleana means that it is a Pteris species with a winged rachis. However, a winged rachis is found in all taxa in the Pteris cadieri complex and is not a useful diagnostic character for this species. In contrast, the monomorphic and regularly bipinnatifid laminae devoid of digital projections is a diagnostic character that distinguishes Pteris grevilleana from all other taxa in the P. caderi complex, except P. dimorpha var. metagrevilleana . As discussed above, these 2 taxa can be distinguished by the width of the sinuses on pinnae (1–2 vs. 3–6 mm).

The cytotypes of P. grevilleana are diploid, triploid, and tetraploid ( Walker 1962; Nakato 1990; Chao et al. 2010, 2012a, b). There are subtle morphological differences among diploid, triploid, and tetraploid P. grevilleana . The main difference is that tetraploid plants are larger (the laminae 8–20 cm long and 8–18 cm wide) than diploids and triploids (both with lamina 3–12 cm long and 4–12 cm wide), and the distance from the sinus to the costule is greater (up to 8 mm in tetraploid vs. less than 4 mm in diploid and triploid). Diploids and triploids are similar in size. Sometimes they can be distinguished by their basal pinnae: straight (diploids) vs. incurved (triploids). Furthermore, diploids and triploids are deciduous in winter, while tetraploids are evergreen ( Chao et al. 2012b). However, these characters, especially the plant size, typically vary with environmental conditions and the maturity of individual plants. The genotypes of the three cytotypes are diploids (X 1 Y), triploids (X 1 *Y) and tetraploids (X 2 X 4 *Y) ( Table 1, Chao et al. 2012b). Because the genotype of tetraploids differs from that of diploids and triploids, cryptic species of indistinguishable morphology within is not separated into different species until new diagnostic characters are found ( Paris et al. 1989).

Plants with a white stripe on the adaxial surface of the pinnae and plants lacking these white stripes are sympatric in China, Taiwan, and Vietnam. Because the plants with different characters are sympatric, the forma rank is preferred to the variety rank . A new combination using the rank of forma is here proposed, i.e., “ Pteris grevilleana Wall. ex J.Agardh forma ornata (Alderw.) Y.S.Chao, H.Y.Liu & W.L.Chiou ”.

Additional specimens examined: — CHINA. Hainan: Jianfengling, Y. S. Chao 1756–1758 ( TAIF) . INDIA. Assam: Lakhimpur Dist., Digbai , March 1889, G. Mann s. n. (K). Namchung: C. B. Clarke 37948c (P) . TAIWAN. New Taipei City: Hsintien Dist. , Mt. Chitang, Y. -H. Chang 20100127-028 –- 031 ( TAIF) ; Wulai Dist. , Mt. Luofeng, P. F. Lu 23294 ( TAIF) . VIETNAM. Thai Nguyen: M. Colani 3442 (P) ; La Flieu, Janvier, 1926, M. Colani s. n. (P). Tonkin: M. l’abbe Bon. 4334 ( SING), 6234 ( HUH, K, P, UC) ; Dam-bs, Lung Wan Village, Sai Wong Mo Shan, W. T. Tsang 29986 ( SING). Vinh Phuc: Tam Dao, P. A. Pételot s.n., 1930 (P) ; Tam Dao Town to Cai Keng Road and Bac Thai Road, Y. S. Chao 1665, 1683–1685, 1701–1703, 1723 ( TAIF, VNMN) .

4. Pteris hainanensis Ching (1959: 166) . Type:— CHINA. Hainan: Gan’en Xian, Chim Fung Shan (Jianfengling), S. K. Lau 5469 (lectotype IBSC-292505!, Figure 7 , here designated; isolectotype MICH-1235048!, Figure 8 ); Pteris cadieri var. hainanensis (Ching) S. H. Wu (1990:54)

Rhizomes short, erect, apex scaly; scales linear lanceolate, 1–2 mm long, 0.2–0.5 mm wide, bicolorous, centers dark brown, margins light brown, entire, apex long-acuminate. Fronds clustered, dimorphic, 15–60 cm long. Sterile fronds 15–40 cm long; stipes green or abaxially dark-red, 8–30 cm long, sparsely scaly at base, grooved on the adaxial side; laminae ovate, 1/3–1/2 length of frond, 8–20 cm long, 5–20 cm wide, pinnate; pinnae linear, 5–16 cm long, 1–5 cm wide, 5 pinnae nearly equally long, simple or with digital projections, apex acuminate to caudate, margins remotely serrate. Fertile fronds 15–60 cm long; stipes 8–30 cm long; laminae pentagonal to ovate, 1/3–1/2 length of frond, 8–30 cm long, 10–20 cm wide, pinnate; basal pinnae with one pair of exaggerated basiscopic pinnules; terminal pinna 1.5–2 times of lateral pinnae, 1–3 pairs lateral pinnae; pinnae linear, simple to with digital projections or pectinate partially, 7–16 cm long, 1–4 cm wide, apex acuminate to caudate; sori along pinna margins; spores tetrahedral if normal or ellipsoidal if aborted, tan.

2 n = 58, 116 ( Chao et al. 2012b).

Ecology and distribution:— China, Malaysia, and Vietnam; in shaded wet places, under evergreen broad leaf forests, below 1,000 m in elevation.

Chinese name:—ẬÑẶŖdz.

Notes:— Pteris hainanensis possesses pinnate laminae and pinnae with digital projections. Some laminae are similar to those of P. cadieri , but those of P. hainanensis are bigger (8–20 cm long and 5–20 cm wide vs. 4–16 cm long and 4–13 cm wide) ( Figure 8 ). Furthermore, the partially pectinate pinnae that result from the many digital projections on fronds of P. hainanensis do not occur in P. cadieri . There are two genotypes in P. hainanensis , but no distinct morphological difference is found between the plants with different genotypes. Therefore a new name is not provided herein until further evidence is found to discriminate those cryptic species ( Paris et al. 1989).

There are two syntype specimens. The one in IBSC is here designated as the lectotype ( Figure 7 ) because it has more mature laminae than the specimen in MICH ( Figure 8 ).

Additional specimen examined: — CHINA. Hainan: W. Y. Chun 6362 ( UC) ; Diaoluoshan, Baishuiling, Y. S. Chao 1259–1263 ( TAIF), Shiqing, Y. S. Chao 1237–1242, 1245, 1246 ( TAIF) ; Jianfengling, W. S. Wu 896-1 ( TAIF) ; Loh-hue M. Mouiuper 230 ( UC) ; Na khi Leng, F. A. McClure 8496 ( US) ; Exianling, Y. S. Chao 1179 ( TAIF) ; Mt. Wuzhi , W. Y. Chun 6659 ( UC), Wuzhishan Fern Survey 271, 607 ( HUH), Shuiman, Y. S. Chao 1225, 1232, 1233 ( TAIF) ; Po-ting, F. C. How 72424 ( HUH). Kwangtung: Renhua, Danxiashan, July 7, 2007, X. Y. Dong s.n. ( TAIF). MALAYSIA. Frareis Hill, B. Molesworth Allen 1189 ( HUH). Pahang: Tembeling, R. E. Holttum 24726 ( UC, US). The Gap, Wu Senangko 8911 (P). VIETNAM. Ba-Ngha : Eberhardt 349 (P). Tonkin : D. A. Billet 4136 (P). Vinh Phuc: Tam Dao Town to Cai Keng Road and Bac Thai Road, Y. S. Chao 1638–1644 ( TAIF) .

5. Pteris incurvata Y.S.Chao, H.Y.Liu & W.L.Chiou , sp. nov. Type:— TAIWAN, New Taipei City, Chungho Dist., Yuhuang Temple, Y. S. Chao 1796 (holotype TAIF-465788!, isotype TAIF-465787!). Figure 9

Evergreen. Rhizomes short, erect, apex scaly; scales linear lanceolate, 1–2 mm long, 0.2–0.5 mm wide, bicolorous, centers dark brown, margins light brown, entire, apex long-acuminate. Fronds clustered, dimorphic. Sterile fronds 9–20 cm long; stipes green or abaxially dark-red, 4–9 cm long, sparsely scaly at base, grooved on the adaxial side; laminae pentagonal to ovate, 1/3–1/2 length of frond, 5–12 cm long, 4–8 cm wide, bipinnatifid; basal pinnae incurved with one pair of exaggerated basiscopic and incurved pinnules, terminal pinna 1.2–1.5 times length of lateral ones; 1(–2) pairs lateral pinnae; rachises winged and pinnae decurrent along distal stipes; pinnae lanceolate, 3–12 cm long, 1.5– 3 cm wide, pectinate; sessile or short-petiolate, apex caudate; margins serrulate; sinuses 1–3 mm wide. Fertile fronds 15–33 cm long; stipes 6–18 cm long; laminae pentagonal to ovate, 1/3–1/2 length of frond, 8–15 cm long, 6–11 cm wide, pinnate; basal pinnae straight or incurved and with one pair of exaggerated basiscopic pinnules, terminal pinna 1.3–2.5 times length of lateral ones; 1–2 pairs lateral pinnae; rachises winged and pinnae decurrent along distal stipes; pinnae linear to lanceolate, 4–20 cm long, 0.5– 3 cm wide, simple to with scattered digital projections or partially pectinate; sinuses 1–3 mm wide; sori along pinna margins; spores tetrahedral if normal, globose or ellipsoidal if aborted, tan.

2 n = 87 ( Chao et al. 2010).

Etymology:— The species is named for its incurved basal pinnae.

Ecology and distribution:— Taiwan, endemic; in shaded wet places, under evergreen broad leaf forests, below 1,000 m in elevation.

Chinese name:—ỆflẶŖdz.

Notes:— The bipinnatifid sterile laminae of this species are wider and shorter (L/W=1.2–1.8) than those of P. dimorpha (L/W=1.5–2.5). The morphology of the pinnae on fertile laminae varies from simple, to having scattered digital projections, to partially pectinate. Both sterile and fertile laminae have incurved basal pinnae.

Additional specimens examined: — TAIWAN. Ilan County: Fushan, Feb . 25, 1998, C. C. Hsu s.n. (TAIF), Y. H. Chang 3054 (TAIF), Y. S. Chao 763, 1152, 1153, 1155 (TAIF). New Taipei City: Chungho Dist., Yuhuang Temple, Y. M. Huang 20050925 -01–10 (TAIF).

6. Pteris perplexa Y.S.Chao, H.Y.Liu & W.L.Chiou , sp. nov. Type:— TAIWAN, New Taipei City, Taoyuan Valley Trail, Dec. 18, 2012, L. Y. Kuo s.n. (holotype TAIF-465789!). Figure 10

Evergreen. Rhizomes short, erect, apex scaly; scales linear lanceolate, 1–2 mm long, 0.2–0.5 mm wide, bicolorous, centers dark brown, margins light brown, entire, apex long-acuminate. Fronds clustered, dimorphic. Sterile fronds 10–20 cm long; stipes green or abaxially dark-red, 4–10 cm long, sparsely scaly at base, grooved on the adaxial side; laminae pentagonal to ovate, 1/3–1/2 length of frond, 5–10 cm long, 4–8 cm wide, pinnate with digital projections to bipinnatifid; basal pinnae slightly incurved and with one pair of exaggerated basiscopic and incurved pinnules; terminal pinna 1.3–2 times length of lateral ones; 1 (–2) pairs lateral pinnae; rachises winged and pinnae decurrent along upper stipes; pinnae linear to lanceolate, 3–10 cm long, 1.5– 3 cm wide, simple to with digital projections to pectinate, sessile or short-petiolate, apex caudate; margins serrulate; sinuses 1–2 mm wide. Fertile fronds 15–40 cm long; stipes 5–20 cm long; laminae pentagonal to ovate, 1/3–1/2 length of frond, 8–20 cm long, 6–12 cm wide, pinnate to irregularly bipinnatifid; 1–2 pairs lateral pinnae, basal pinnae straight or slightly incurved and with one pair of exaggerated basiscopic pinnules; terminal pinna 1.3–1.8 times length of lateral ones; rachises winged and pinnae decurrent along distal stipes; pinnae linear to lanceolate, 4–20 cm long, 0.5– 3 cm wide, simple to with digital projections or pectinate partially; margins serrulate; sinuses 1–3 mm wide; sori along pinna margins; spores tetrahedral if normal, globose or ellipsoidal if aborted, tan.

2 n =87 ( Chao et al. 2010).

Etymology:— The species is named for its great morphological variation, which is greater than that of any other species in the P. cadieri complex.

Ecology and distribution:— Japan and Taiwan; in shaded wet places, under evergreen broad leaf forests, below 1,000 m in elevation.

Chinese name:—ṘAẶŖdz.

Notes:— This species and two similar species in the P. cadieri complex, possess bipinnatifid sterile laminae and pinnate fertile laminae. Pteris dimorpha is diploid and has straight basal pinnae, whereas P. perplexa and P. incurvata are triploids and have incurved basal pinnae.

The sterile laminae of P. perplexa are regularly bipinnatifid or occasionally pinnate with digital projections, whereas the fertile laminae vary from pinnae with digital projections to partially pectinate. The incurved, sterile basal pinnae are similar to those of P. incurvata , but their curvature is not as great. The different genotypes (PgiC) indicate these two cryptic species arose from different hybrid events and have different parental lineages ( P. perplexa X 3 *Y vs. P. incurvata X 2 X 3 Y; Chao et al. 2012b). Because their morphologies are different even if subtle, P. perplexa and P. incurvata are designed here to provide multipurpose usages for the herbarium curator, conservation biologist, park naturalist, etc. ( Paris et al. 1989).

This species is found only in Taiwan and Japan. In the Pteris cadieri complex in Japan, only this species and P. grevilleana have been found.

Additional specimens examined: — JAPAN. Kagoshima: Tokunoshima Isl., Mt. Amagi-sam, M. Tagawa & K. Iwatsuki 2580 ( KYO). Okinawa: Ishigaki Isl., Mt. Omoto-dake, S. Tawada 77 ×4 ( KYO) . TAIWAN. Ilan County: Suao Township, Suhua Ancient Trail , Nov. 5, 2009, W. H. Wu s.n. ( TAIF), Y. H. Chang 20081121-007, -008 ( TAIF). New Taipei City: Taiping Village, Y. S. Chao 732, 733, 1112 ( TAIF) ; Neitung National Forest Recreational Area, Y. S. Chao 1366 ( TAIF) ; Pinglin Dist. , Mt. Chiuchiungken, P. F. Lu 17726 ( TAIF), Jan. 3, 2009, P. F. Lu s.n. ( TAIF) ; Shuangsi Dist., P. F. Lu 2775 ( TAIF), Shihsun Village, P. F. Lu 5140 ( TAIF) ; Sijhih Dist., Hsinshan-menghu, P. F. Lu 778, 779, 7860 ( TAIF), Y. H. Chang 4066, 4067 ( TAIF), H. L. Chiang 2279 ( TAIF) ; Taoyuan Valley Trail , Y. S. Chao 745, 749 ,

767, 1415 (TAIF), P. F. Lu 24809 (TAIF). Taoyuan County: Longtan, Shihmenshan, S. C. Liu 700 (TAIF). Taipei City: Yangmingshan, Y. S. Chao 1518 (TAIF).

Figures

FIGURE 6. Lectotype of Pteris grevilleana Wall. ex J.Agardh in E (Wallich 2680).

FIGURE 7. Lectotype of Pteris hainanensis Ching in IBSC (S. K. Lau 5469), a mature plant with only fertile laminae.

FIGURE 8. Isolectotype of Pteris hainanensis Ching in MICH (S. K. Lau 5469), a juvenile plant with both sterile and fertile laminae.

FIGURE 9. Illustration of Pteris incurvata Y.S.Chao, H.Y.Liu & W.L.Chiou, sp. nov. A, based on the holotype (Y. S. Chao 1796); B & C, based on two laminae from the isotype (Y. S. Chao 1796).

FIGURE 10. Illustration of Pteris perplexa Y.S.Chao, H.Y.Liu & W.L.Chiou, sp. nov. Based on the holotype (L. Y. Kuo s.n.).