World Botanical Associates Web Page
Prepared by Richard W. Spjut
April 2003; Photos added May 2006; Jan. 2007, Mar 2007, June 2007
Last updated Aug 2007, May 2014
|Fig. 11. Comparison of leaves of extant T. mairei (clear photos, isolectotype, P) with extinct T. engelhardtii (grainy photos, reproduced from Kvaček 1986), from an Oligocene deposit in Bohemia.|
14. Taxus mairei (Lemée & Lév.) S. Y. Hu ex T. S. Liu (Figs. 1C, 3B, 11, 63–67, 225, 227), Illustr. Native & Introd. Lign. Pl. Taiwan 1: 16. 1960. Tsuga mairei Lemée & Léveillé, Monde des-Pl. sér. 2, 16: 20.1914. Taxus chinensis Rehder var. mairei (Lemée & Lév.) W. C. Cheng & L. K. Fu, Fl. Hupehensis 1: 28. 1976 (also indicated as comb. nov. in Fl. Reipub. Pop. Sin. 7: 443. 1978). T. wallichiana var. mairei (Lemée & Lév.) L. K. Fu & Nan Li, Novon 7: 264. 1997. Type: China. Yunnan, Dongchuan, 700–800 m, May 1912, Maire s.n., “holotype” indicated by Rehder (1936) at E (isotype: fragment also at A!). Other duplicate or original material distributed by E at BM! (annotated by Spjut Oct. 1997 as T. mairei var. speciosa [Florin] Spjut ined.) and at P (Fig. 11, also shown here).
14a. var. mairei. Maire yew. Distribution: Forest margins, 300–1300 m; China (Sichuan, Yunnan, Anhui, Guizhou, Guangdong, Guangxi(?), Jiangxi, Fujian, Hunan, Zhejiang, Taiwan).
Tree 5–20 m high; branchlets equally divided , the main branch showing in a zigzag pattern, yellowish-green, reddish to orange with age; bud scales mairei not persistent, or withered in age, in 3 obscure ranks, yellowish-brown, minute deltoid, thick, closely adpressed, upper scales carinate near apex, 0.3–0.5 mm long. Leaves spreading at nearly right angles to branchlets, adjacent leaves closely parallel, oblong (or slightly ovate to ellitpical in type), machetelike, arcuate near base and upturned near apex, unevenly tapered to an acute apex, 1.5–2.5 cm long, 3–4 mm wide, 200–350 µm thick, dark glossy green and slightly convex above to acute midrib that is channeled along base, midrib more obtuse above the mid region towards apex, paler and duller green and concave below to a squarrose elevated midrib that is shallowly channeled, often lustrous reddish-brown in herbarium specimens; upper (adaxial) epidermal cells in transverse section wider than tall, 10–12 µm tall, 25–37 µm wide, ellipitical in transverse section, thin-walled; lower epidermal cells larger than upper on midrib and marginal regions, 15–30 µm tall, 20–40 µm wide, numbering 12–30 from the margin to stomata band, 18 or less on the midrib, abruptly differentiated from those in stomata band, irregularly squarrose to rectangular, most 1–3× l/w near margins and on midrib, occasionally 5–10×l/w on midrib, inflated (mamillose), papillose on up to nearly half of the cells between margin and stomata band, occasionally papillose on outer midrib cells; papillae variable in shape and size, positioned medially in stomata band and adjacent marginal cells, submarginally on outer midrib cells; stomata bands pale orange, or yellowish, broader than marginal border; stomata in 12–16 irregular continuous rows/band, often crowded, stoma sometimes with a blackish halo. Male cones scales 4-seriate in bud, not seen at maturity. Female cone subcylindric, 4 mm long in bud, scales 6-seriate, basal one conduplicate, all greenish, aril forming a shallow cup, red at maturity; seed purplish, ovoid, 3–4 mm long, 3 mm wide.
Representative Specimens—China—Sichuan: Pei-pah, tree, Law 65 (K). Yunnan: N Yunnan, Tie’tchang-keou, rares, rocheres des forêts, 700 m, Maire 19B (P), Yunnan without data, Maire (BM); Yung Chun, 6 May 1913, Rankin s.n. (K). Anhui (Anhwei): Southern, Chanen, 300 ft, in woods by grave-yard, rare, tree 60 ft, 1.8 ft dbh, R-C. Ching 3168 (A, K). Guizhou: Jiangkou Xian, Daiyenpeng along the Kaitu River on the SW side of the Fanjing Shan mountain range, 750-1000 m, tree 18 m, Sino-Amer. Guizhou Bot. Exped. 1046 (A, BM). Guangdong (Kwangtung): Loh Ch'ang District, Chong Uen Shan near Kau Fung, dry sandy soil, in thicket, fairly common, W.T. Tsang 20694 (A, BH, K, NA, S, US). Jiangxi (Kiangsi): NW, Si-ho, Hwang-kong-shan Mt, tree, common on slope, Y.K. Hsiung 6443 (A); Ningdu, Y?ntungtsch?, Wang-Te-Hui 445 (A); Lienhwa-shan, 800 m, Wang-Te-Hui 458 (A). Hunan: Ma-Ling-Tung, Sinning Hsien, 600 m, tree 30 ft, aril yellowish becoming red, edible, C.S. Fan & Y.Y. Li 664 (A). Zhejiang (Chekiang): Taishun Hsien, open place by bridge, tree 25 m, Y.L. Keng 317 (A); Tien-Tai-Shan, 1300 m, tree 5 m, 3–4 in dbh, shady woods, H.H. Hu 342 (A, US); same locality, C.Y. Chiao 14510 (A, US); same locality, Cheng 3617 (US); without specific locality, S. Chen 1044 (A). Fujian (Fukien): Yeuping, Shih-Sun-Keng village, on hill, 650 m, tree 4 m, H.H. Chung 2865 (A, BM, K); Buong Kang, mountain slope back of village, 700 m, H.H. Chung 3581 (A); western, 1868-1873, David (P). Taiwan: Hualien #1,2,4,5,7,8,9,10, TD3, C-j Chang, 1 of 3, 19 Jan 1993; 2-of-2, 8 Feb 1993 (no additional data), C-j. Chang (wba, collections in bold; determination to variety depends of branching character, branchlets supplied by Chang were insufficient to evaluate this feature, identification of var. speciosa for collections not in bold are based on variable shape in epidermal cells of young leaves, and the leaves remaining mostly green when dried).
Taxus mairei is distinguished from other species of the Sumatrana Group by the two ranked leaves spreading at nearly at right angles, by the abaxial surface of leaves having a truncated but elevated midrib with epidermal cells larger in diameter than those on the adaxial surface (as seen in T-sect.). The midrib is usually channeled, especially in the mid region of the leaf.
In annotations I have recognized two varieties by differences in branching and leaf arrangement; one has isodichotomous ± zigzag branching with dark reddish-green leaves that spread closely parallel to one another; the other—corresponding to the type of T. speciosa (e.g., H. Smith 10398 from Sichuan; Plate 5 in Florin 1948a)—has monopodial branching with more greenish colored leaves arranged in a less parallel manner. However, the syntypes of T. mairei in further study were found to correspond mostly to an intermediate complex as previously described under T. celebica—one that appears closer to T. speciosa than to T. mairei. Nonetheless, the morphological differences seem to have taxonomic merit, and one solution is to recognize a third variety or nothotaxon.
The difference between var. mairei and var. speciosa is is further supported by differences in seed shape and seed color, but this is based on the paucity of herbarium material with seed, while in other species seed traits can vary on individual plants. Generally, specimens of T. mairei with monopodial branching and greenish leaves have tan colored, globose, glossy seeds (widest above the base) with a distinct brownish apical nipple, whereas specimens with reddish leaves—spreading more closely parallel to one other—have purplish glazed seeds without a brownish nipple. In comparison to other species, seeds of T. kingstonii in specimens from China are similarly shaped to those of the latter variety but also differ in being slightly smaller, duller, and more varied in color—from tan to purplish, or in a specimen from India, seed was angled instead of rounded to apex; such angled seeds are characteristic of the T. cuspidata complex. Seed was rare in other related species; in one specimen of T. celebica from Vietnam seed is conical as in T. chinensis and purplish colored as in T. mairei, but since other specimens from Vietnam (e.g., Soulie s.n.) could be hybrids between T. wallichiana and T. celebica, I cannot attach much taxonomic value to seed characters in the one specimen from Vietnam.
Taxus mairei has a long history of nomenclatural confusion. The original authors thought they had a species of hemlock (Tsuga). Rehder (1936), upon discovering that Tsuga mairei Lemée & H. Léveillé (Léveillé 1914) belonged to Taxus, treated it as a synonym of T. chinensis; however, the ICBN (Art. 11.4) requires the earlier epithet, mairei, be adopted (S. Y. Hu, in Liu, Illustr. Nat. Ind. Lign. Pl. Taiwan 16. 1960). Nevertheless, Cheng & Fu (1978), in maintaining this illegitimate name, made another illegitimate combination—T. chinensis var. mairei—that they earlier introduced invalidly—without reference to the basionym (Cheng et al., 1975; Art. 33.2, Greuter et al., 2000); it has since been transferred—T. wallichiana var. mairei (Lemée & Léveillé) Fu & Li (in Li & Fu 1997)—without accounting for the type of T. baccata var. sinensis Henry (in Elwee & Henry, Trees Gr. Brit. & Irel 1: 100. 1906).
Regardless of the nomenclatural misapplications of the name, the taxonomic application in recent years has been to treat yews in subtropical China under the epithet “mairei” if their leaves lack papillae on the abaxial midrib. The epithet sumatrana, an earlier available name, was not applied because the Chinese botanists had not seen its type (Cheng & Fu 1978; Hu 1964; Li & Fu 1997); instead, they selected mairei for the varietal epithet—placing it under T. wallichiana.
14b. Taxus mairei var. speciosa (Florin) Spjut, J. Bot. Res. Inst. Texas 1(1): 240. 2007. Taxus speciosa Florin, Act. Hort. Berg. 14, 8: 382 (1948). Type: China, Guizhou, [Fanjing Mts.] Kiangkow, 450 m, in light woods, tree 8 m high, fruit red—8 Dec 1930, Y. Tsiang 7525— holotype S (isotypes: A! BM! K! NY [photocopy!] US! Fig. 231).
Special yew. Distribution: Forest margins, near streams or open areas on hillsides; 100-750 (-1600) m; China (S Shaanxi, Sichuan, NE Yunnan, Guizhou, N Guangxi, Hunan, Guangdong, W Hubei, Jiangxi, Zhejiang, Fujian).
Similar to T. mairei. Branchlets mostly unequally divided, not in a zigzag pattern, yellowish-green turning pale orange with age. Leaves more unevenly spaced along one side of branchlets, overlapping slightly at their base in pairs, not closely parallel along margins, machete-like or nearly linear, arcuate (falcate) near base and upturned near apex, unevenly tapered to an obtuse or acute apex, to 3.5 cm long, 3–4 mm wide, light green and convex above to acute midrib that is channeled along base, midrib more obtuse above the mid region towards apex, paler and duller green and concave below to a squarrose elevated midrib that is shallowly channeled, often greenish and puckered in herbarium specimens with a rugose adaxial (dorsal) surface; stomata in (9-) 11–18 (-21) irregular rows/band. Male cones developing from most leaf axils on the current branchlets, scales 4-seriate. Female cone subcylindric, 4 mm long in bud, scales 6-seriate, basal one conduplicate, all greenish, aril forming a deep cup, red or yellow at maturity; seed often tan in color, ovoid with a nipple at apex, 3–4 mm long, 5–6 mm long or diam.
Representative Specimens—China—Ningxia Huizu: Sikong, Lung Dung An, 10 km from city, 1000 m, C.Y. Chiao 1223 (GH, S). China—Shaanxi: Mt. Tsin-lin to Lao-lin, 3000 m, large tree. Sichuan: western, Nin Ya-chou Fu, 2000 ft, tree 30–5 ft x 8-15 ft, Wilson 1265 (A, BM, K, S, US); S of Kuan-hsien, ridge, 1390 m, tree 30 ft, 3 in dbh, F.T. Wang 20600 (A); S of Kuan Hsien, ravine, 1160 m, tree 15 ft, 2 in dbh, F.T. Wang 20541 (A); Kuan-hsien, Mt. Tsing-cheng, Chengtu and vicinity, shady places in forest, common, tree 8 m, W.P. Fang 12205 (A, BM); Tienchuan Hsien, Tienchuanchow, 2500–3000 ft, in woods, tree 30 m, 4 ft dbh, W.P. Fang 3461 (A, K, P); same locality, tree 25 m, 3 ft dbh, W.P. Fang 3442 (A, K, P, US); Kuan Hsien, Chien-Chang-Shan, 1000 m, temple yard, tree 40 ft, male cones yellow, C.S. Fan & Class 91 (A); Shikong [Zigong?], Tien-Chuan Ling-Kwan, 3000 ft, back of old temple, erect tree 50 m, S.Y. Hu 1563 (A); Nanchuan Hsien, roadside, tree 10 m, W.P. Fang 5811 (A, K, P); Ta(s)chienlu, Cheng 1001 (BM), Cheng 1475 (P); Li County, Jiabigon, 2500 m, in forest, tree 5-7 m high, Z. Quing-sheng 0152 (BM); W Oua, Pao Shan, NE, 600 m, Legendre 586 (P); W region, between Huangnipu and Yaan (Yachou), Malingtsang, 900 m, H. Smith 10402 (A, BM, S); E Sichuan: Tchen-Keou-Tin, Farges 128 (P); Yun-Ling-Hsien, 5,000 ft, roadside, tree 15 m, 2 ft dbh, W.P. Fang 3796 (A). Yunnan: Forrest 15053 (K). Zhejiang (Chekiang): Yen Tang Shan, C.Y. Chiao 14618 (A, K, US); without locality data, S. Chen 1063 (A); S. Jentang, hillside near stream, tree 30 m, 2 ft dbh, H.H. Hu 97 (A); Siachu, open cultivated hilltop, 2600 ft, rare, tree 50 ft, 1.2 ft dbh, R-C. Ching 1676, with papillose midrib; S. Chekiang, open moist hillside, rare, tree 60 ft, 1.8 ft dbh, R-C Ching 2489 (A, K, US). Guizhou (Kweichow): Pichish, along roadside, tree 7 m, Y. Tsiang 8987 (A, P); Liang Feng Yah, Tsunyi Hsien, 900 m, tree 10 m, 20 cm dbh, Steward et al. 154 (A, BM, S); Songtao Xian, vicinity of Lengjiaba in the vicinity of the confluence of the Xiaohe and Dahe rivers, NE side of the Fanjing Shan mountain range, 820-1120 m, forest margin, tree 4 m, Sino-Amer. Guizhou Exped. 1981 (GH). Guangxi/Guizhou (border): 15 mi S of Bin lon(g) Hiu Shan W Inchen, 4000 ft, small tree, 2 ft, fairly common in woods, Ching 5976 (A, US). Jiangxi: Lu Shan, along stream in forest, Chiao 18795 (US). Hunan: Changning Hsien, Yang-Shan, 680 m, on slope in forest, tree 40 ft, C.S. Fan & Y.Y. Li 296 (BM, GH). Hubei (Hupeh): Metasequoia Area, between Ta-yin-pin & Chunglo, in valley, vicinity of Shui-sa-pa, 900 m, tree 15 m, 18 cm dbh, Gressitt 2507 (GH); vicinity of Li-chuan, on open hill, 3600 ft, C.T. Hwa 164 (A). Guangdong (Kwangtung): Hung-mio to Mio-lan, near Jui-feng, Lokohong Hsien, N.R. Region, 1340 m, front of temple yard by stream, tree 50 ft, 16 in dbh, aril red, seed green, Tsiang Ying 1425 (A, P); Ruyuan Xian, Nanling Exped. 1838 (A). Guizhou (Kweichow): Ta Ho Yen, Kianakou Hsien, rocky slope near farm house, 980 m, tree 10 m, 35 cm dbh, Steward et al. 328 (A, BM, K, P, S, US). Fujian (Fukien): Naping, 800 m, dense woods, Guo-sheng 1544 (K, US); Ing-dan E. Fookma, Price 1258b (K). Taiwan: C-j Chang, Hualien #1,2,4,5,7,8,9,10 (wba, identifications based on color and broad abaxial leaf margins; branching is generally required to distinguish var. mairei from var. speciosa).
Taxus mairei var. speciosa has been recognized by branchlets arising anisodichotomously, and by leaves appearing greenish and unequally spaced along branchlets in dried specimens (Spjut 2007b). It occurs more inland whereas var. mairei is found more along coastal provinces; however, exceptions include the type from N Yunnan and related specimens in Sichuan that have the isodichotomous branching of var. mairei and greenish colored leaves of var. speciosa. Variety speciosa is further distinguished by leaves often sickle-shaped, uniformly curved, or curved more abruptly well above the base, or if not curved then tapering from near mid region of the leaf as opposed to near base in T. celebica, or near the apex in var. mairei. The exception is the type specimen of var. mairei that differs in leaves appearing widest near the base as in T. celebica but still has leaves closely parallel as in seen in all other specimens included.
Taxus speciosa was described by Florin to distinguish a yew in China based on leaves that had a smooth midrib from one other in China that he recognized to have a papillose midrib, which he had determined to represent T. wallichiana var. chinensis based on study of its type. The taxonomic feature of a smooth midrib was also erroneously described for T. chinensis by Handel-Mazzetti (1929). Florin (1948) correctly realized that T. chinensis was closely related to T. wallichiana even though he only mentioned the type for T. chinensis and not T. wallichiana, although he had studied an isosyntype. However, Florin incorrectly provided another name for the species he distinguished by the absence of leaf papillae on the abaxial midrib; he cited not only an earlier name—Cephalotaxus celebica, but also its type (Warburg 16889). Apparently, elements in the protologue did not conform to his concept of T. speciosa. Other authorities soon realized this error (Cheng & Fu 1978), and Florin's name was replaced by T. mairei S. Y. Hu (Liu 1960), who had also distinguished it from T. chinensis according to Florin taxonomy as evident by her 1955 annotations of Harvard specimens (Hu 1964). Notwithstanding, this name was soon replaced by an earlier name T. celebica Li (1963) who recognized only one species in China even though Hu (1964) indicated she adopted T. mairei because the type for T. celebica might differ based on one specimen she saw from Sulawesi that had a papillose midrib. She might have further assumed that the Berlin type of T. sumatrana was destroyed; however, Florin took a fragment of the Berlin type to Stockholm. It is surprising that an earlier name mentioned by Pilger (1903)—C. sumatrana, had been neglected—until finally applied by de Laubenfels (1978). Although T. speciosa was founded upon nomenclatural error, it can still be recognized if its type can be shown to belong a distinct species. Nevertheless, Möller et al (2013) recognized a new species based on the absence of papillae on the abaxial midrib; the morphological key is dé·jà vu, Florin (1948).
This variety has also been difficult to define because its leaves vary widely in shape and epidermal features, especially young leaves. On the abaxial surface, epidermal cells vary from rectangular as in T. celebica to trapezoidal as in var. mairei. Stomata bands may be found with anywhere from (8-) 11–21 rows/band, and the development of papillae on adjacent epidermal cells seems to vary according to age of the leaf.