Taxus umbraculifera

24. Taxus umbraculifera (Siebold ex Endl.) [Ravenscroft] [C.] Lawson, Abietineae—List Pl. Fir Tribe No. 10, 80 (1851). Cephalotaxus umbraculifera Siebold ex Endlicher, Syn. Conif. 239 (1847); Taxus cuspidata Siebold & Zuccarini var. umbraculifera Makino, Illus. Fl. Nippon, 910 (1931), Fig. 2730. Taxus cuspidata var. ambraculifera Makino, Illus. Fl. Nippon (1925). Original material unknown. Type: Japan. Neotype (designated by Spjut 2007)—illustration in Makino, Illus. Fl. Nippon, Fig. 2730 (see Fig. 153). 1931; epitype (designated by Spjut 2007)—Japan, Honshu: Hyogo Pref., Mt. Hyonosen, Muroi 30, ex Herb. Hiroshi Muroi, Kobe, identified by Muroi as T. cuspidata var. ambraculifera Makino (A!).

Shrub or tree with erect to ascending, numerous, short, thick branches, 1–10 m or more high; branchlets subverticillate to alternate, crowded toward ends of branchlets, dull gray or brownish with orange tint; bud-scales persistent at base of current growth, paleaceous, pale to dark brown, or metallic greenish-brown, 2–3 seriate, thick, turgid, the lower scales more acutely folded along midrib to form a keel, the uppermost scale concave, ca. 1–1.5 mm long. Leaves mostly green in the herbarium, in equidistant spirals from sharply reflexed (not twisted) petioles, the blades often turned sideways, rather than facing upwards as in T. caespitosa, almost in a decussate arrangement in some plants, crisscrossing more than overlapping when pressed, oblong, 1–2 cm long, 1–3 mm wide, 300–350 µm thick, dull dark green and broadly convex above to a rounded midrib that forms a shallow channel at base along the upper surface, pale yellowish-green and concave below to a rounded midrib, plane to recurved ca. 45° near margins; upper (adaxial) epidermal cells in transverse section elliptical, 10–20 µm tall, 25–40 µm wide, lower (abaxial) nonstomatal cells similar, 10–12 µm tall, 25–40 µm wide, numbering 10–15 (-22) between margin and stomata band, the marginal border of cells abruptly differentiated from those in the stomata band, uniform in width, most 1–3× l/w in the marginal region, 3–5× l/w on midrib, epappillose up to 8–10 (-15) cells from margins, occasionally papillose on midrib; stomata bands broader than the marginal region, (8-) 9–11 (-14) rows (-17 rows in var. hicksii). Male cone scales 5-seriate, sharply angled in bud, ca. 3 mm in diam. at maturity; pollen sacks ca. 8. Seed cones maturing on current season growth, scales 4–5 seriate; seed nearly globose to short conical, 4 mm long, 3–4 mm diam.

The origin of the name T. umbraculifera may be in horticulture (cf. Gordon 1858, 1875); however, Siebold had recognized Cephalotaxus umbraculifera, according to Endlicher (1847) who provided a detailed description, but who also expressed doubt as to whether the species was distinct from Taxus cuspidata. Ravenscroft (Lawson et al. 1851) attributed C. umbraculifera to Siebold and Zuccarini, and indicated it was a synonym of T. cuspidata. The epithet suggests an umbrella-like leaf arrangement as shown above, but the leaves were described as mostly two-ranked (Endlicher (1847). Ravenscroft (Lawson et al. 1851) indicated that branches are verticillate with “distichous” branchlets. I have not seen any original material, and herbaria that I have contacted in this regard for Siebold specimens reportedly have none. Nonetheless, an illustration in Makino (1931) clearly depicts the umbrelliform leaf arrangement that is distinctive for this species. In later manuals on the flora of Japan (e.g., Ohwi 1965), var. nana Rehder (1902) became the name for this taxon, but this is antedated by var. microcarpa Trautvetter (1859).

The leaves of T. umbraculifera appear in star-like (almost decussate) manner when looking down the branch from apex, and unlike T. caespitosa, the blades of T. umbraculifera will face different directions. Leaves appear perpendicular to the plane along one side of a branchlet, and in the same plane of along another side of branchlet. This is not easily determined in pressed specimens; however, the crisscrossed blades partly reflect this, which is in contrast to the radial orientation of leaves in T. caespitosa that all twist in the same manner, appearing imbricate when looking down the apex of the branchlet. The leaves of T. umbraculifera are also sharply reflexed at their petioles in contrast to bending upwards (erect) in T. caespitosa; this difference accounts for the two-ranked appearance in T. umbraculifera in contrast to the secund appearance in T. caespitosa, often seen on older branchlets.

In Europe, a similar type of leaf arrangement occurs in T. recurvata. These taxa may be linked through T. contorta var. mucronata, and forms of T. chinensis, which exhibit crisscrossing leaf arrangements near apex of branchlets (e.g., Henry 7097 from Sichuan). The taxonomic significance of these traits may have become lost through introgression with other species.

Four varieties of T. umbraculifera are recognized according to apparent differences in habit and leaf arrangement. Variety umbraculifera, variable in habit, but mostly arborescent with wide spreading branches, has leaves appearing radial on erect branchlets, especially near apex, and decussate to nearly two-ranked on horizontal branchlets. Variety hicksii is distinctly columnar with ascending branchlets and mostly radial leaves; var. microcarpa, also variable in habit, is distinguished by isodichotomous branching and by leaves that slightly overlap and crisscross and not particularly secund near apex of branchlets, and var. nana that is mostly a low shrub with densely crowded branchlets and leaves.

24a. Var. umbraculifera (Figs. 153, 177–179). Shrubs or trees. Branchlets ascending or drooping; leaves mostly radial near apex, dull dark green when dried, oblong.

Umbrelliform yew. Distribution. Japan, Manchuria.

Representative Specimens. Japan: Honshu: Mt. Ooyhama, Kanagawa-Pr, cult., 25 Oct 1952, shrub 2 m high, Suzuki 499003 (A); Shunane Pref., Mt. Sentsu-zan, Naito s.n. (A). Manchuria (Mandshuria) SE: Ex herb. hort. bot. Petro. yr 1860, Maximowicz (S). Mandshuria SE, ex herb. hort. bot. Petro. yr 1860, Maximowicz (P: 2 specimens on one sheet).

Japan—Honshu: Hyogo Pref., Mt. Hyonosen, Muroi 30, epitype (A). Illustration attached to specimen indicates abaxial leaf margin lacks papillae across 8 cells followed by 4 rows of papillose cells, 10 stomata rows, and midrib of 10 thin-walled, yellowish, epapillose, epidermal cells.	Japan—Honshu: Shunane Pref., Mt. Sentsu-zan, Naito s.n. (A). Illustration attached to specimen indicates abaxial leaf margin lacks papillae entirely across 8 cells. This is followed by 11 stomata rows in a yellowish green band, and a midrib of 14 yellowish, epapillose cells.	Japan—Hokkaido: Sapporo, Arimoto s.n., June 1903. Illustration attached to specimen indicates abaxial leaf margin lacks papillae across 13 cells. This is followed by 2 rows of papillose cells, 11 stomata rows, and a partially papillose midrib of 15 cells, papillae lacking in the center 5 rows of midrib cells. Leaf x-section shows a double palisade layer, the second row of cells being much smaller than the primary row.
Japan—Honshu: locality illegible, Wilson s.n (A). Illustration attached to specimen indicates abaxial leaf margin lacks papillae entirely across 13 thin-walled epidermal cells. This is followed by 11 stomata rows in a greenish stomata band, and midrib of 13 greenish, epapillose epidermal cells. The leaf x-section shows a double palisade layer of cells, both palisade rows of equal length. Of additional note is the determinations of subsp. globosa f. tardiva on one label and cuspidata var. nana by Wilson on his label.	Japan—Honshu: Nippon, forests and in cultivation, collector unknown (A). Illustration attached to specimen indicates abaxial leaf margin lacks papillae across 13 thin-walled epidermal cells. This is followed by 4 rows of papillose cells, 11–13 stomata rows in a pale yellowish stomata band, and midrib of 11 pale yellowish, epapillose epidermal cells. The leaf x-section shows a double palisade layer of cells, both palisade rows of equal length. Photo on right shows male cones and cuspidate scales at base of branchlets.	Japan: Honshu: Mt. Ooyhama, Kanagawa-Pr, cult., 25 Oct 1952, shrub 2 m high, Suzuki 499003 (A). Illustration attached to specimen indicates abaxial leaf margin lacks papillae entirely across 15 thin-walled epidermal cells. This is followed by 14 stomata rows in a greenish stomata band, and papillose midrib of 17 cells, the papillae low. The leaf x-section shows a double palisade layer of cells, both palisade rows of nearly equal length.
Mandshuria SE, ex herb. hort. bot. Petro. yr 1860, Maximowicz (P: 2 specimens on one sheet). Illustration indicates abaxial leaf margin lacks papillae entirely across 10 epidermal cells. This is followed by 12 stomata rows, and midrib of 15 papillose epidermal cells. The leaf x-section shows a single palisade layer of cells.	Mandshuria SE, ex herb. hort. bot. Petro. yr 1860, Maximowicz (S). Illustration indicates abaxial leaf margin lacks papillae across 8 epidermal cells. This is followed by 6 rows of papillose cells, 12 stomata rows, and a papillose midrib. The leaf x-section shows a single palisade layer of cells.	This is similar to the Wilson specimen above. Collection data unclear, N. Koshimotowke s.n., yr 1910. (US: 1311889). Illustration indicates abaxial leaf margin lacks papillae entirely across 9 epidermal cells. This is followed by 12 stomata rows and a yellowish epapillose midrib of 18 cells. The leaf x-section shows a single palisade layer of cells.
Top Left: Sheet in Gray Herbarium (GH) with three specimens. In the upper left corner is a specimen from Lugd Batav, which compares favorably with a duplicate specimen at the Paris Museum of Natural History (P) shown directly below it. The specimen on the right in the top left photo is from a male plant, accompanied by a label from Herb. Zuccarini, reportedly collected by de Siebold in 1842. A close-up view of the specimen is shown in the top right photo. This would appear to be original material, one of many uncited specimens employed by Siebold & Zuccarini in regard to selecting a type for Taxus cuspidata. There is also a small branchlet with seeds accompanied by a handwritten notation from Blake that the seed branchlet came from Yeso, which historically refers to the island of Hokkaido, collected what appears to be in 1861. The lectotype was chosen from the original material at M. The specimen in GH, however, is not a duplicate or isolectotype. It is regarded as T. umbraculifera var. microcarpa. The seed bearing branchlet is also this variety. This determination is supported by the relatively small cones, wide branching, and leaves not notably secund near apex of branchlets. The specimens from Herb. Lugd. Batav. are referred to T. umbraculifera var. umbraculifera.	China—JiLin, JiAn. JA001 Identification features: Sharply reflexed crisscrossed leaves, generally shorter near apex in contrast to those of T. biternata

24b. Taxus umbraculifera var. hicksii (Rehder) Spjut (Figs. 172–173), J. Bot. Res. Inst. Texas 1(1): 278. 2007. Taxus cuspidata var. hicksii (hort.) Rehder ex Bailey, Cult. Evergreens 189 (1923). Taxus media Rehder f. hicksii, J. Arn. Arb.: 198 (T. cuspidata hicksii Hort., synon.). (1923). Type—Hort.: Hicks Nursery, Wesbury, Long Island, NY, A 8036, holotype (Fig. 173, A!).

Shrub, cylindrical-columnar in outline but not tapering as in T. fastigiata; branches mostly erect, brownish to maroon, not reddish-orange; branchlets arising from a distinct erect leader. Persistent bud-scales conspicuous, 5-seriate, pale brown. Leaves crowded, radial on young shoots, crisscrossing when pressed, two-ranked to slightly radial on older shoots, linear, averaging ca 1.5 cm. long, 1.5–2.0 mm wide, dark green and convex above to a rounded midrib, yellowish and nearly plane below to rounded midrib, notably curled in dried specimens; abaxial epidermal cells usually partly papillose between stomata bands and margin, the papillae positioned marginally; midrib without papillae, or papillose more on the outer 2-3 rows of cells,, or entirely with low papillae; stomata 11-17 rows/band. Seed angular or smooth, ovoid, ca. 4–5 mm long.

Hicks yew. Distribution: Japan

This is distinguished by the erect to ascending branches and branchlets with linear leaves that spread in whorl like arrangements, and by leaves with papillae along the abaxial marginal zone. It differs from the typical variety by its columnar habit. The leaves may appear two-ranked on lower branches, but if these branches are turned over, the underside of the branch usually has some of the leaves reflexed. It differs from T. caespitosa by the persistent bud-scales, and by the revolute leaf margins in dried specimens.

This horticultural variety appears to occur naturally in Japan based on six specimens from there, but it may have independently evolved there as a hybrid, or it may have been introduced into Japan from North America since the specimens cited above were collected during the mid 1950's—after the Hicks yew was described by Rehder (1923). Moreover, the Hick's yew reportedly originated from seed of “T. cuspidata ‘Nana’” sometime around 1900 (den Ouden & Booom 1965), which has been generally interpreted to include native plants in Japan characterized by radial leaves (Ohwi 1965). The cultivars and apparent wild plants share the darker color foliage on older branchlets, but differ slightly in having a less papillose marginal zone on the abaxial surface of leaves. Other cultivars may be included under this variety for classification purposes even though they are undoubtedly of hybrid origin. One example is a broadly rounded bush with relatively short branchlets and leaves arranged in nearly in two-ranks that appears as a hybrid between the Hicks yew and T. biternata. Others that I have seen at the Secrest Arboretum could be hybrids between T. fastigiata and T. biternata.

Representative Specimens: Japan. Iwate-Pref.: Asagishi, Muroi 3593 (A). Nagano Pref.: Kamikochi, Muroi 3715 (A). Gifu- Pref.: Takayama, Muroi 3698 (A). Hyogo Pref.: Kumatugi, Mikata-gun, Muroi 5603 (A); Mt. Hatibuse Muroi 5424 (A); Wakasugi, Muroi 5648 (A). Cultivation: private residences in MD, 8495 Imperial Drive, Laurel; 757 Dunberry, Arnold.

Cultivated: Type (A)	Japan. Iwate-Pref.: Asagishi, Muroi 3593 (A).. Illustration indicates abaxial leaf surface lacks papillae across 12 marginal cells followed by 8 rows of papillose cells, 12 stomata rows, and a midrib 15 cells wide, the outer two rows papillose.	Japan. Nagano Pref.: Kamikochi, Muroi 3715 (A). Illustration indicates abaxial leaf surface lacks papillae across 14 yellowish marginal cells followed by 2 rows of papillose cells, 11 stomata rows, and a midrib 14 cells yellowish cells without papillae.
Japan. Hyogo Pref.: Kumatugi, Mikata-gun, Muroi 5603 (A). Abaxial leaf surface lacks papillae across 13 marginal cells followed by 11 rows of papillose cells, 9-10 stomata rows, and a midrib 15 cells wide, the outer three rows obscurely papillose	Japan. Wakasugi, Muroi 5648 (A). Illustration indicates abaxial leaf surface lacks papillae across 10 marginal cells followed by 16–17 stomata rows, and a midrib 14 cells wide, all with low marginal papillae. Although the marginal region lacks a transitional zone of papillose cells, this would seem to be offset by the higher count of stomata rows.	Japan. Gifu-Pref.: Takayama, Muroi 3698 (A). Illustration indicates abaxial leaf surface lacks papillae across 13 marginal cells followed by 2 rows of papillose cells, 11-12 stomata rows, and a midrib 12 cells wide.
Japan. Mt. Hatibuse Muroi 5424 (A). Illustration indicates abaxial leaf surface lacks papillae across 8 marginal cells followed by 6 rows of papillose cells, 12 stomata rows, and a midrib 15 cells wide, the outer three rows obscurely papillose.

24c. Taxus umbraculifera var. microcarpa (Trautv.) Spjut (Figs. 156–171, 283–288), J. Bot. Res. Inst. Texas 1(1): 279. 2007. Taxus cuspidata Siebold & Zucc. var. microcarpa (Trautv.) Kolesnikov, Bull. Far E. Branch Acad. Sci., USSR 13: 43, Fig. 2. 1935. Taxus baccata L. var. microcarpa Trautvetter in Maxim., Mem. Acad. Sci. St. Petersb. Sav. Etrang. 9: 259 (Prim. Fl. Amur.). 1859. Type not specified; original material (syntypes) from several locations, one from Manchuria (1 May 1855, reportedly sterile, GH! P!), and another from Sakhalin Is. (Weyrich, Sep. 1853, with fruit, A?). Type—Exped. Soc Geogr., 1855 (1 May), Manchuria [China], Heilongjiang, Maack s.n., Lectotype: lower of two specimens at GH!; Isolectotype: upper of two specimens at P!

Taxus umbraculifera ssp. laxa Spjut ined. Type: Korea. Ooryong- too (Oagelet Island), 0-900 m, bush or small tree, common—Wilson 8538, holotype at A! (isotype at US!).

Shrubs or trees, branchlets isodichotomous, or subpinnate; bud-scales persistent on current season growth, 2–3 seriate, mostly small, 0.5–1.5 mm long, deltoid, thick, concave, smooth or with a thickened midnerve above, the lower scales nearly ovate and abruptly acute. Leaves mostly dark green in the herbarium, spreading radially on young shoots, overlapping more than crisscrossing when pressed, mostly two-ranked below, oblong, 1.4–2 cm long, 2–3 mm wide, 275–400 µm thick, ca. 8× l/w, dull olivaceous and broadly convex above to a rounded midrib, yellowish green and more strongly concave below to a flush to slightly rounded midrib, plane to recurved ca. 45º near margins; lower (abaxial) epidermal cells numbering 14–20 between margin and stomata band, the marginal cells gradually differentiated from those in the stomata band, pentagonal to subquadrate near margins, 1–3 (-7)× l/w in the marginal region, (1-) 3–6× l/w on midrib, mostly epappillose, or occasionally papillose on 4 outer rows of midrib cells and on 1–5 rows in the marginal zone nearest the stomata band; papillae 1–3 rows across each cell; stomata 7–14 rows/band, with blackish halo. Male cone scales overlapping in 3 ranks, pollen sacks 5–6. Seed angular or rounded, conical, 3–5 mm long, 3–4 mm diam, yellowish or purplish.

Small seed yew. Distribution: Common, NE temperate Asia.

Representative Specimens. Russian Federation—Primorie [Primorskiy] Prov., vicinity of Vladivostok, Palczevsky 3601 (A, K, US); Primorie Prov., Bay of Peter, the First Sea Reserve, Island of Stenin, 26 Apr 1979, Kypehinova s.n. (in Russian, A); Geogr. Exped. 1855, Maack (P). Manchuria Region: Korea septentrionalis *provincia Pen-nian Muorum Jahn...Frajectus Lpatan-ien, 27 Jun 1897, Komaróv 88 (A); Rossica, Aultzo Prov., Ussuri [Ussuzieusis] insula Afnold, Komaróv 88 (P); Manchuria: Rossica Palczevsky [Komaróv] 88 (BM, K). Korea: Nemon-rei, Kyongsan, Kyogsan Prov., tree up to 50 ft, common, 12 Oct 1917, Wilson 9332 (A); Kyaraboken, cult., annotated var. nana on Wilson label s.n., annotated f. tardiva ex herb. 2–4-1914, Sakurai s.n. (A); Herb. Lugd. Batv. (P); Zuccarini 593, in adnot. T. baccata (M). Japan—Hokkaido: Ishikasi Prov., Apr 1884, K. Muijabe s.n. (A). Nanokwa, Tosa, 18 Apr 1888, Watanabe s.n. (A); Sapporo, Agric. College, 15 Jun 1885 (A), Jun 1878 (A); Sapporo, Siebold, ex Herb. Zuccarini (GH); Mt. Nantai, Lake Chuzenji, 20 Aug 1904, Mochizuki s.n. (A). Honshu: Yokohama, yr 1862, ex Herb. USDA 1888 (US: top specimen); Japan, no locality data (US: 1311889); Hida, Takayama, 17 Sep 1910 (A); Yokohama, yr 1862, ex Herb. USDA 1888 (P: p.p.; US: lower of 2 specimens); Kiaraboku s.n. (US: 1311889).

Japan—Hokkaido: Sapporo, Agric. College, 15 Jun 1885 (A). Illustration attached to specimen indicates abaxial leaf margin lacks papillae across 13 cells in which there are several rows of quadrate cells nearest margin, followed by 3 rows of ± rectangular cells and 3 rows of inflated cells. The marginal cells also include 3 rows of papillose cells. This is followed by 9–10 rows of stomata, and 12 midrib cells, which also lack papillae. The leaf in x-section is shown to consist of a single palisade layer. Note small seeds.	Japan—Hokkaido: Sapporo, Agric. College, June 1878 (A). Illustration attached to specimen indicates abaxial leaf margin lacks papillae entirely across 12 cells in which there are several rows of quadrate cells nearest margin, followed by 3 rows of inflated cells. This is followed by 10 (-12) rows of stomata, and midrib cells with low papillae. Note small scattered male cones.	Korea: Nemon-rei, Kyongsan, Kyogsan Prov., tree up to 50 ft, common, 12 Oct 1917, Wilson 9332 (A). Illustration attached to specimen shows abaxial leaf margin lacks papillae entirely across 10 cells in which there are several rows of quadrate cells nearest margin, and 3 rows of inflated cells near stomata band. This is followed by 12 rows of stomata in a greenish band, and 17 midrib cells. papillae developing on outer rows. Leaf in x-section shows a single layer of palisade cells.
Korea: Ooryong- too (Oagelet Island), 0-900 m, bush or small tree, common—Wilson 8538 (A). Illustration attached to specimen shows abaxial leaf margin lacks papillae entirely across 12 cells in which there are 3 rows of inflated cells near stomata band. This is followed by 12 rows of stomata in a greenish band, and a midrib with papillae developing on outer rows. Leaf in x-section shows a double palisade layer. Additional illustration based on specimen in US herbarium. It notes there are 17 midrib cells, none are indicated to be papillose, while the remaining features are the same as the specimen in A.	Manchuria: Heilongjiang, Exped. Soc Geogr., 1855 (1 May), Maack s.n., Isolectotype (P).	Japan—Hokkaido: Sapporo, Agric. College, April 1884 (A). Illustration attached to specimen indicates abaxial leaf margin lacks papillae across 14 cells in which there are several rows of quadrate cells nearest margin, followed by 2 or more rows of inflated cells. This is followed by 12–14 rows of stomata, and 17 midrib cells, which lack papillae. The leaf in x-section is shown to consist of a single palisade layer. Note similarity of specimen to type from Manchuria.
Russian Federation: Primorie Prov., Bay of Peter, the First Sea Reserve, Island of Stenin, 26 Apr 1979, Kypehinova s.n. (in Russian, A). Illustration attached to specimen shows abaxial leaf margin lacks papillae entirely across 5 cells in which there are 3 rows of inflated cells near stomata band. This is indicated to be followed by 4–5 rows of papillose cells (papillae submarginal), ±10 rows of stomata, and a epapillose midrib of 14 cells. Leaf in x-section shows a double palisade layer.	Russian Federation—Primorie [Primorskiy] Prov., vicinity of Vladivostok, Palczevsky 3601 (K). Illustration shows abaxial leaf margin lacks papillae entirely across 8 cells. This is indicated to be followed by 11 rows of stomata, and a epapillose midrib of 15 cells. Leaf in x-section shows a double palisade layer. Leaf material was evidently not a good specimen for sectioning as the illustration is somewhat incomplete. A more complete illustration is shown in the specimen from A, and noted are 4 rows of papillose cells bordering the stomata band. Of additional note is the determination by Keen, T. cuspidata cv. 'Densa' (K) .	Honshu: Yokohama, yr 1862, lower specimen, Maximowicz, ex Herb. herb. horti. bot. Petropolitani; upper specimen USDA 1888 (P: p.p.; US, pp). Duplicates of this arrangement are found in different herbaria, sometimes the Maximowicz specimen is on top, in other cases it is mounted below. The collection of the 2 specimens supposedly is to show mature male and female cones of Taxus cuspidata, assuming of course, it is the only species in the Sino-Japanese region. Thus, one may never really know for sure just where these specimens were collected even though reported from “Yokahama.” The top specimen has branching characteristics of T. umbraculifera var. microcarpa, but the relatively large male cones in aggregate compare more favorably with var. umbraculifera. The illustration indicates an the abaxial leaf near margin lacks papillae across 7 cells, and is followed by 4 rows of papillose cells. Stomata band has 10 rows of stomata. Midrib was noted to be similar in color to the stomata band, and of 12 papillose cells, the center 5 rows appearing less papillose. The parenchyma cells were noted to be golden brown in color, and the palisade is shown to be a double layer of cells. The lower specimen includes several seeds near the apex of a branchlet, the seed is wider than tall; this agrees with T. umbraculifera var. microcarpa.
K BM P P Manchuria Region: Korea septentrionalis provincia Pen-nian Muorum Jahn...Frajectus Lpatan-ien, 27 Jun 1897, Komaróv 88 (A, BM, K, P). Illustrations show abaxial leaf margin is 14-18 cells wide, lacking in papillae entirely, or with 4 rows of papillae cells, the epidermal cells nearest the margin are smaller, generally with 5–9 rows of cells, and those nearest the stomata band without papillae are inflated in 2–5 rows. This is followed by a stomata band with 12–14 rows of stomata, and an epapillose midrib, measured in one specimen as 12 cells wide. Leaf in x-section shows a double palisade layer in one specimen and a single layer in two specimens.

Taxus umbraculifera var. microcarpa is recognized primarily by isodichotomous branching, oblong leaf shape (ca. 8x l/w) and phyllotaxy in which leaves spread in slight crisscross arrangement and are not particularly secund near apex. These features are not easily recognized without study of many specimens.

Trautvetter originally distinguished var. microcarpa from T. baccata by the smaller seed (Maximowicz 1859). Kolesnikov (1935) further indicated it was a rounded shrub 0.5–1.5 m high and 5–7 m in diam. that reproduced by layering, which he illustrated. The two yews he recognized (tree and shrub yews) are parapatric in SE Russia, occurring in different ecological habitat. I have recently concluded that there is wide diversity of shrub forms of Taxus in SE Russia, Korea, and Japan, some of which I include under var. microcarpa, while others I consider new varieties. One example, a specimen from Manchuria collected by Komaróv (P), differs by the divaricate branching and presence of what appears to be also a layering branch, which has leaves similar to that of var. T. canadensis var. adpressa. Duplicates at other herbaria lack the layering branch but are apparently from the same or similar plants. A specimen collected by Palczevsky (3601) at K from the SE Russian coast near Vladivostok, annotated “Taxus cuspidata cv. ‘Densa’” by Keen (July 1966), appears more densely branched.

Collector's notes on herbarium labels such as by E. Wilson indicate this can be a large bush or tree, which in further studies may warrant taxonomic distinction.

24d. Taxus umbraculifera var. nana (Hort. ex Rehder) Spjut (174–176), J. Bot. Res. Inst. Texas 1(1): 281. 2007. Taxus cuspidata var. nana Hort. ex Rehder in Bailey, Cyclopedia Amer. Hort. 1773 (1902). Taxus cuspidata f. nana (Rehder) Wilson, Conif. Taxads Japan 13 (1916). Described from horticulture, no specimens cited. Original material unclear. Neotype—Japan: Honshu: Pref. Hyogo, Mt. Hyonosen, 1100–1400 m, in Fagus forest with sasa thicket, on ridge, shrub 2 m, fr red, 11 Aug 1983, Murata 44671, det. as T. cuspidata var. nana Rehder (A!).

Low shrub, densely branched, leaves mostly radial, crisscrossing, oblong, dark glossy green above, paler beneath.

Dwarf yew. Distribution: E Russia (islands), Japan.

The epithet “nana” implies a dwarf plant, and Rehder (1902) described T. cuspidata var. nana as a “dwarf compact form with shorter leaves” in regard to a horticultural plant. Rehder (1949) later considered it only a form. Nevertheless, others applied the varietal name to native yews in Japan. They were characterized as a low shrub with radial leaves occurring mostly along the sea-side of Japan (Ohwi 1965). I distinguished this variety from others by the radial reflexed leaves as indicated above.

Representative Specimens—Russia: Far East: Primorskiy Region, Sea Reserve, Island Bolshoy Pelis, Borzova s.n. (in Russian, A); Sakhalin, ex herbario horti Petropolitani, Augustinowiez, T. baccata var. microcarpa, Schmidt, p.p. with T. caespitosa var. caespitosa (A). China: Shanxi (“Schenhsi merid”): Taipei-schan, 1936, G. Fenzel 972 (A). Japan: Mt. Fujiwara Mie pref., Muroi 1969 (A); Mt. Himekami, Iwate pref, 14 Sep 1955, Muroi 5933 (A); Japan, no additinal data, Faurie s.n. (P). Cultivars. Secrest Arboretum, Ohio: ‘Newport,’ ‘Hatfield.’

Honshu: Pref. Hyogo, Mt. Hyonosen, 1100–1400 m, in Fagus forest with sasa thicket, on ridge, shrub 2 m, fr red, 11 Aug 1983, Murata 44671, det. as T. cuspidata var. nana Rehder (A!), proposed neotype.	Japan: Mt. Himekami, Iwate pref, 14 Sep 1955, Muroi 5933 (A). Illustration attached to specimen indicates abaxial leaf margin lacks papillae entirely across 10 epidermal cells. This is followed by 10–12 stomata rows in a greenish stomata band, and midrib of 10 epapillose epidermal cells. The leaf x-section shows a double palisade layer of cells, both palisade rows of equal length.	Russian Federation: Far East: Primorskiy Region, Sea Reserve, Island Bolshoy Pelis, Borzova s.n. (in Russian, A). Illustration attached to specimen indicates abaxial leaf margin lacks papillae entirely. This is followed by 14 stomata rows stomata band, and a partially papillose midrib of 18 epidermal cells, the outer 4 rows with marginal papillae. The leaf x-section shows a double palisade layer of cells, both palisade rows of equal length.
Japan: Mt. Fujiwara Mie pref., Muroi 1969 (A). Illustration attached to specimen indicates abaxial leaf margin lacks papillae entirely across 14 epidermal cells. This is followed by 12–13 stomata rows in a greenish stomata band, and midrib of 20 epapillose epidermal cells. The leaf x-section shows a double palisade layer of cells, both palisade rows of equal length.	China: Shanxi (“Schenhsi merid”): Taipei-schan, 1936, G. Fenzel 972 (A).	Cultivar: Secrest Arboretum, OH, Newport Media, 31-264. Branchlets with relatively small obtuse, bud-scales. papillae lacking across 8 marginal cells followed by 4 obscurely papillose cells; stomata in 10-11 rows; midrib smooth, 22 cells wide; palisade double. These features agree with T. umbraculifera var. nana. The broadly 4-lobed seed near apex may be seen as evidence of a hybrid with a variety of T. baccata.