Taxus caespitosa

21. Taxus caespitosa Nakai, Ch[Ty]ôsen Sanrin Kaihô (J. Kor. For. Soc.) 158: 40 (1938). Taxus cuspidata Siebold & Zucc. var. caespitosa (Nakai) Q. L. Wang, Clavis Pl. Chinae Bor.-Or., ed. 2: 73 (1995). Type—Japan: Honshu, Tottori Pref., Mt. Daisen. Nakai cited two specimens from the type locality: “Arika Kimura, Aug. 1924.—typus; Taketaro Sawada, July 15, 1922.” LECTOTYPE (designated in Spjut 2007b): Sawada s.n., 15 July 1922 (TI!). Topotypes: Wilson (A!), Makino 43792 (S!).

Taxus umbraculifera ssp. latifolia (Pilger) Spjut ined. (in adnot. A)

Plants prostrate to erect shrubs, mostly open places, on mountain-tops, ridges, and slopes; branches and leaves spreading uniformly, or often to one side—with a windswept appearance (secund). Leaves all erect, or some erect, often radial on erect branchlets, secund in part or entirely on spreading to recurved branchlets, usually thick, strongly resinous, oblong, obtuse, more convex above than concave below, thickened and rounded along margins, often revolute along margins when dried; epidermal cells elliptical in transverse section, 8–12 µm tall, 10–25µm wide. Seeds often small, pale yellow to purplish in color (probably stained by the aril), conical like a ‘Hershey Kiss.’

Taxus caespitosa is allied to T. cuspidata by leaf anatomy, but more similar to T. umbraculifera in phyllotaxy. Differences in leaf arrangement between T. caespitosa and T. umbraculifera appear related to the habit of the plant and the habitat type to which each has adapted. Taxus caespitosa has erect leaves that appear radial on erect branchlets and secund on horizontal branchlets. This is obviously an adaptation to growing in sun-exposed habitats as leaves of most yews will show this phototropic response (secund leaves), and plants in such habitats are mostly shrubs. Taxus umbraculifera, by contrast, has sharply reflexed leaves—at their petiole junction, and while they also appear radial on young erect shoots, they exhibit a different phototropic response by staying reflexed regardless of the branch orientation. The leaves in T. umbraculifera often appear distichous when a horizontal branch is viewed from above, but when looking down the branch from apex, the leaves appear to spread in two or more decussate ranks. The erect to ascending branchlets along with the manner in which leaves spread are features obviously associated with a tree (habit) growing in thickets or woodlands.

21a. Var. caespitosa (Figs. 154–158). Low rounded bushes, branches prostrate with aerial roots, layering, branchlets erect to ascending, fastigiate to alternately arranged, or sympodial, densely compacted, 1–2 m high; bud-scales 3–4 seriate, longest scales 1.5–2 mm long. Leaves often becoming reddish in the herbarium, ascending to erect in whorl-like arrangements, densely crowded on branchlets, closely overlapping, oblong, 1–2 (-2.5) cm long, 2–4 mm wide, 350–500 µm thick, glossy olivaceous and broadly convex above to a rounded midrib, below yellowish-green and slightly concave to a rounded midrib, recurved slightly (ca. 15º) near thickened margins, with a ventral (abaxial) marginal region of 8–12 (-18) smooth cells wide, the marginal cells abruptly differentiated by size from those of the stomata band, uniform in width, 1–3× l/w, 3–5× l/w on midrib, lacking papillae, or papillae evident on up to 4 rows of cells, papillose or epapillose on midrib; papillae positioned submarginally on midrib cells to medially on accessory cells in stomata bands, 1–2 rows across each cell; stomata in 9–11 rows/band. Male cone scales 4-seriate, sharply angled in bud, 3–4 mm in diam. at maturity; sporangiophores flattened, with thick ribs, slightly dilated near base, pollen sacks ca. 8. Seed rounded, appearing more conical than in other varieties, widest near base, 3 mm long, 3–4 mm diam., pale yellowish to purplish, flattened at base to the attachment point, tapering to the apex from the middle to a short pale nipple.

Caespitose yew. Distribution: Russia (Sakhalin Is.), Korea, Japan.

Taxus caespitosa var. caespitosa is recognized by densely congested branchlets and radial leaves that appear more imbricate than decussate with an erect orientation by curving upwards along petioles and also along blades in the same direction they spiral. Wilson (1916) described plants he saw in Japan as having prostrate branches (from which erect branchlets apparently arise). This is evident in herbarium specimens by the one-sided development of branchlets, which includes specimens from the type locality, Mt. Daisen, obtained by Wilson and by Makino. However, the cultivars referred to this variety are not prostrate shrubs, and do not show the small seeds as seen in specimens of wild plants.

In E Asia where shrub yews apparently have diversified in habit and phyllotaxy, T. caespitosa allegedly exemplifies an ancestral type from which others such as var. latifolia may have been derived. The one sided development of branchlets and leaves appear as ancestral traits (of var. caespitosa) retained in the erect branches of var. latifolia. Kolesnikov (1935) regarded the presence of yew in the Manchurian flora as an archaic element having little historical relationship to the rest of the vascular flora, while shrub yews in other regions have been noted to occur in environments that are distinct from their tree relatives such as reported by Elias (1992) for yews he studied in Ukraine and Georgia. I have also recognized a distinct shrub variety in the Pacific NW (T. brevifolia var. reptaneta), and in NE America T. canadensis has long been know for its shrubby monoecious habit.

Other yews with a phyllotaxy similar to T. caespitosa occur in Morocco, southern Spain, and on the island of Sicily. They are relatively rare and show a closer relationship to T. baccata by the persistent reddish, obtuse bud-scales, and by leaves appearing papillose across the abaxial surface to four cells from the margin. It is interesting that yew fossils from Europe are more like extant species in E Asia in leaf anatomy. Perhaps, the Morocco yew, the most distinct among these, retained ancestral feature of radial leaves, while acquiring leaf anatomical traits through introgression with species such as T. contorta that allegedly entered the European flora from northern Asia during the late Tertiary. Related yew specimens from Portugal and southern Spain (Spanish yew) have slightly longer leaves in less definite whorls, and they in turn are related to another variety in Algeria and France (French yew) that has more laxly arranged leaves. These alleged relicts occur near the southwestern-most distributional range of the genus in the Euro-Mediterranean with another alleged relict, T. canadensis, as seen in Morocco, southern Spain, and on Madeira.

Russia—Sakhalin Is.: Schmidt s.n. (GH). Korea: in forest, Aug 1907, Faurie 1512 (A, BM, E). Japan—Honshu: Mt. Daisen, 2000 m, [forms a dense scrub] dense shrub, 1–2 m [covering large areas on the wind-swept middle and upper slopes], 30 Nov. 1914, Wilson s.n. (A) [specimen given to Wilson, Wilson 1916]; Mt. Daisen, Makino 43792; Niigata, Yuzawa-machi, Minamiuonuma-gun, in Pinus pumilascrub, 1650 m, evergreen shrub 0.8 m high, fr red, 3 Oct 1979, Taoda 3887 (A); Honshu, Hakkada, bush 1–1.5 m, windswept slopes, 1000–2000 m, rare, 4 Jul 1914, Wilson 7133 (A); Honshu, Mutsu Prov., Mt. Hakkoda, Mizushima 1985 (A); Ohobu near Kobe Calta, 19 Mar 1955, Muroi 1058 annotated T. cuspidata var. ambraculifera (A). Cultivars—Ohio, Secrest Arboretum: ‘Colean’, A30-131; ‘Dwarf Japanese Yew’ A30-163; ‘Hill Anglo-Japanese Yew’ A30-264.

Honshu: Mt. Daisen, Makino 43792 (topotype, S). Illustration indicates abaxial leaf surface lacks papillae across 8 marginal cells across followed by 4 rows of papillose cells, 10–11 stomata rows per band and a midrib 12 cells wide appearing mamillose.

Honshu: Mt. Daisen, Wilson s.n. (A, topotype). Illustration attached to specimen indicates abaxial leaf surface lacks papillae across 8–9 marginal cells followed by 4 rows of papillose cells, 10–11 stomata rows per band and a midrib 11 cells wide

Russia—Sakhalin Is.: Schmidt s.n. (GH), with T. umbraculifera var. microcarpa. Illustration indicates abaxial leaf surface lacks papillae across 11 cells, stomata are in 11 rows, and midrib is 16 cells wide, the outer 2 rows with papillae.

Honshu, Hakkada, bush 1–1.5 m, windswept slopes, 1000–2000 m, rare, 4 Jul 1914, Wilson 7133 (A). Illustration attached to specimen shows abaxial leaf lacking papillae across 8 marginal cells, 11 stomata rows per band, and midrib 11 cells wide.

Honshu: Niigata, Yuzawa-machi, Minamiuonuma-gun, 1650 m, evergreen shrub 0.8 m high, fr red, Taoda 3887 (A).

Honshu: Mutsu Prov., Mt. Hakkoda, Mizushima 1985 (A). Illustration attached to specimen indicates abaxial leaf surface has 18 marginal cells (no papillae), 11 stomata tows, and midrib is 11 cells wide (no papillae).

Korea: in forest, Aug 1907, Faurie 1512 (A). Illustration from specimen at BM, shows abaxial leaf margin to lack papillae across 9 marginal cells, stomata 9-10 rows, and a papillose midrib.

Honshu: Ohobu near Kobe Calta, 19 Mar 1955, Muroi 1058 (A).

Cultivars: Secrest Arboretum, Wooster, OH.

21b. Taxus caespitosa Nakai var. angustifolia Spjut (Figs. 163–164, 271), J. Bot. Res. Inst. Texas 1(1): 268. 2007. Type: Japan. Prov. Kozuke, Oze-ga-hara, “shrubby habit, “leaves dark green,” “yellowish green beneath,” 26 Aug 1950, Mizushima 401 (Figs. 163–164 (holotype A, leaf with a double row of palisade parenchyma cells, lacking papillae across 14 marginal cells, with 9–10 stomata rows/band, seed).

Apparently prostrate bearing numerous erect reddish orange branchlets, persistent bud-scales few, cuspidate, ~ 1 mm. long; leaves mostly distichous, spreading from erect branchlets, erect on horizontal branchlets, reportedly dark green above, yellowish below, becoming reddish green in the herbarium, revolute along margins when dried, 1.5–2.0 mm long, ~1.5 mm wide, lacking papillae entirely across 14 marginal cells and on and midrib; with 9–10 stomata rows/band. Seed near base of branchlets, ovoid, purplish, tapering to a sharp apex.

Ground cover yew. Korea, Japan.

Text Box: Fig. 271: Taxus caespitosa var. angustifolia, reproduced from Illustrated Encyclopedia of Fauna & Flora of Korea; Vol. 5 (1965), as T. caespitosa. This variety is recognized by the relatively thin, narrow leaves as in T. biternata, but apparently has the habit of T. caespitosa var. caespitosa as reproduced here from the Illustrated Encyclopedia of Fauna & Flora of Korea; Vol. 5 (1965). This reference reported this species to reproduces by layering.

21c. Taxus caespitosa Nakai var. latifolia (Pilger) Spjut, J. Bot. Res. Inst. Texas 1(1): 269. 2007 (Figs. 159–162, 273–275). Taxus cuspidata var. latifolia Nakai, J. Kor. For. Soc. 158: 39 (1938). Taxus baccata L. [ssp. cuspidata (Siebold & Zuccarini) Pilger] var. latifolia Pilger, Pflanzenreich 4(5): 112 (1903). Taxus cuspidata var. latifolia Nakai, J. Kor. For. Soc. 158: 39 (1938). Syntypes from E Russia, Korea and Japan. Lectotype (designated by Spjut 2007)—Japan: Faurie 6345 at P! (with two labels, one dated 30 Dec. 1890 and another dated 1888, with male cones); isolectotype at K! (fragment).

Taxus media Rehder f. hatfieldii Rehder, J. Arnold. Arb.: 198. 1923. Taxus media var. hatfieldii Wilson, Horticulture, n ser. 3: 30, fig. (1925). Type: from horticulture (holotype A!).

Broad-leaved yew. Distribution. E Russia, Korea, Japan.

Shrubs or trees, variable in habit and branching; main stems prostrate to many spreading upwards from a common base. Branchlets often recurved. Leaves radial on erect branchlets, secund on lateral branchlets, often greenish orange in the herbarium; abaxial marginal cells of (6-) 10–18 smooth cells across, usually with several more quadrate rows of cells near margins, the cells enlarged or rectangular in several or more rows towards stomata bands, 1–3× l/w, 3–5× l/w on midrib, with papillae evident on up to 6 rows of cells, papillose or epapillose on midrib; papillae positioned submarginally in 2 rows on each cell; stomata in 7–13 (-16) rows/band. Seed conical, 3 mm long, 3–4 mm diam., pale yellowish to purplish.

Representative Specimens—Russia Federation—Sakhalin Is.: 46º37'N, 142º53'E, Prigarodne, mixed conifer (Abies, Picea)/broadleaved (Betula, Sorbus) woodland, 150 m, to 1.8 m high, Flanakan &Kirkham 203 (K). Korea: in forest, 800 m, 19 Jul 1910, Taquet 4455 (A); without locality data, Faurie 117 (P), 3406 (P), 5975 (P). Japan—Hokkaido: Shiribeshi Prov., Shiribeshi-san, branches wide spreading, 1–2 m, common, 1300-2000 m, 27 Jul 1914, Wilson 7265 (A). Honshu: Mutsu Prov., Mt. Hakkoda, erect tree, 8 Jul 1952, Mizushima 1989 (A). without locality data, Faurie 5114 (P: 2 sheets). Locality unknown: Folley s.n., identified T. cuspidata ‘nana’ (K).

Variants—Russia—Sakhalin Is.: Dvorakovskia & Bokina s.n. (A). “Mandshuria” SE: ex herb. hort. bot. Petro. yr 1860, Maximowicz (P [Bunge], S:C-2114). Japan—Honshu: Kanagawa Pref., Mt. Imaizumi, Tomitar ex. Makino 43780 (S: C-2120); Honshu, Akita Pref. Makino 43769 (S); Nikko, Shimotsake, Apr 1887 (P); Nikko, May 1888 Shimotsake 446 (US). Hokkaido: Kitami Prov., Aug. 1897, Sapporo Agric. College (PH)

Russia Federation—Sakhalin Is.: 46º37'N, 142º53'E, Prigarodne, mixed conifer (Abies, Picea)/broadleaved (Betula, Sorbus) woodland, 150 m, to 1.8 m high, Flanakan &Kirkham 203 (K). Illustration shows leaf x-section and cell shapes on abaxial surface. The abaxial leaf margin is indicated to lack papillae across 15 cells. This is further shown to be divided into 3 rows of small irregularly quadrate cells nearest the margin, followed by 3 rows of somewhat rectangular cells, 4 rows of inflated cells and 5 rows of more regular rectangular cells nearest the stomata band. The stomata band is indicated to have only 8 rows of stomata. The subsidiary cells were noted to be orange, from which it may be deduced that the specimen was kept in cold conditions before being pressed. Papillae are shown to be low and marginal on accessory cells, and alternate on outer 14 midrib cells. The palisade is a double layer of cells, a feature that is common to the Cuspidata Alliance.

Korea: Faurie s.n. yr 1910 (A). Stomata rows 13.

Japan: Faurie 5114, yr 1889 (P). Illustration shows leaf abaxial margin to lack papillae across 6 cells, three rows of which are inflated, followed by 4 rows of papillose cells, and 12 rows of stomata. Midrib is noted to be 15 cells wide with obscure medial papillae.

Korea: Faurie 3406 (P). Illustration shows abaxial leaf surface to lack papillae across 10 marginal cells of which 3 rows nearest the margin appear relatively small and irregularly quadrate, followed by 7 rows of inflated cells, and 6 rows of papillose cells. The stomata are noted to be in 13 rows. The abaxial midrib is noted to be 14 cells across with low submarginal papillae. The leaf x-section indicates a marginal count of 11 cells across without papillae; thus, the abaxial leaf marginal zone without papillae might be indicated as 10-11 cells wide.

Japan: Nikko, Schimotsake, yr 1880 (P). Annotated T. cuspidata in Oct 1997, but now considered T. caespitosa var. latifolia by the narrow margin of 8 cells, and short epapillose cells on abaxial midrib.

Japan? Faurie 5975 (P). Illustration shows abaxial leaf margin to lack papillae across 10 cells in which 1-2 rows nearest margin are relatively small; this is followed by 7 rows of papillose cells and 7 rows of stomata. Midrib is shown to lack papillae.

Cultivated. from Hungary, Folley s.n., identified as Taxus cuspidata nana (K). Illustration indicates abaxial leaf epidermis lacks papillae across 16-17 cells, 1-2 rows nearest margin are relatively small, followed by 2 rows of slightly larger irregularly quadrate cells followed by 13 rows of irregular long hexangular to somewhat rectangular shaped cells, stomata rows are indicated to be 13; midrib cells is shown to lack papillae. The palisade layer is indicated to be double.

Japan? Faurie 5714 (P). Illustration shows abaxial leaf surface to lack papillae across 14 marginal cells in which two rows nearest margin are short and irregularly quadrate, followed by 2 rows of slightly larger cells, followed by 10 rows of much longer cells; stomata band has 13 stomata rows' midrib has medial papillae.

Cultivar: Halorian yew, Secrest Arboretum OH, A30-116. Leaf 2.3 mm wide. Abaxial leaf margin lacks papillae across 16 marginal cells in different size groups (4 + 9 + 3); stomata 12 rows, midrib 12 cells wide, smooth.

Cultivar: Colean yew, Secrest Arboretum OH, A30-131. Abaxial leaf margin lacks papillae across 8 marginal cells; stomata dense in 11 irregular rows, midrib 15 cells wide, papillose.

Erect with Spreading or Ascending Branches

Japan: Faurie 6345, yr 1888. Lectotype (P). Illustration notes abaxial leaf surface lacks papillae across 18 marginal cells, 3 nearest margin are much smaller; there are 11 rows of stomata and the midrib, which is 18 cells wide, lacks papillae. Leaf x-section shows elliptical epidermal cells and double palisade layer, the second rows of cells shorter than the first. Photo on top right shows close-up of male cones; lower right shows stomata band, 250x.

Japan, mountains, Faurie 117, yr 1888 (P). Illustration shows abaxial leaf surface to lack papillae across 8 marginal cells, followed by 4 rows of longer obscurely papillose cells, 8 rows of stomata and a papillose midrib 18 cells wide. The parenchyma cells in the leaf mesophyll differ from what is commonly seen.

Honshu: Mutsu Prov., Mt. Hakkoda, erect tree, Mizushima 1989 (A). Illustration attached to specimen indicates abaxial leaf surface to lack papillae across 8 marginal cells and also across the entire midrib, which is 16 cells wide. Stomata are noted to occur in 9–10 rows.

Japan—Hokkaido: Shiribeshi Prov., Shiribeshi-san, branches wide spreading, 1–2 m, common, 1300-2000 m, Wilson 7265 (A). Illustration attached to specimen indicates abaxial leaf surface lacks papillae across 16 marginal cells with two rows of shorter cells nearest margin, and also across the entire midrib, which is 12 cells wide. Stomata are noted to be in 11–12 rows, and surrounded by yellowish orange subsidiary cells. Leaf x-section shows a double palisade layer.

Honshu: Nikko, May 1888 Shimotsake 446 (US). Leaf sections from this specimen show a strong resemblance to T. mairei var. speciosa; however, the leaf has a thickened lip-like margin as evident in x-section. The abaxial leaf surface lacks papillae across 16 marginal cells in which 5 rows of cells nearest the margin were narrower and longer; stomata were noted to in 15 rows and the abaxial midrib has enlarged epidermal cells lacking papillae entirely, 15 cells wide.

Hokkaido: Rishiri, Sapporo Agricultural College, yr 1888 (PH). Abaxial leaf margin was noted to be 11 cells across without papillae, stomata in 11 rows and midrib of 14 cells wide was papillose on outer 4 rows.

Korea: Mt. Halla, close-up of midrib cells, 250x

Intermediates to Taxus baccata and T. umbraculifera

Russian Federation—Sakhalin Is.: Dvorakovskia & Bokina s.n. (A). Illustration attached to specimen indicates abaxial leaf surface lacks papillae across 8 marginal cells, followed by 14 rows of papillose cells, 11 rows of stomata, and the midrib 11 cells wide. Leaf in x-section shows a double palisade layer. This specimen was annotated T. fastigiata based on color and leaf arrangement, although these features and also flexuous branchlets compare closely with T. baccata var. glauca. However the double palisade layer, 8 marginal cells, and seed tapering from near base are features that favor classification in the Cuspidata Alliance.

Honshu: Akita Pref. Makino 43769 (S). Illustration indicates abaxial leaf margin lacks papillae across 7–8 marginal cells; stomata bands are yellowish orange with 12 rows of stomata, and midrib lacks papillae. This specimen has branching similar to T. baccata var. jacksonii whereas the color on leaves and branchlets is close to var. glauca. Also, bud-scales are rather inconspicuous as in T. baccata. However, the relatively wide leaf margin and stomata count support placement in the Cuspidata Alliance.

Japan—Honshu: Kanagawa Pref., Mt. Imaizumi, Tomitar ex. Makino 43780 (S: C-2120). The long branches suggest T. cuspidata, whereas whorled branching and thick branches suggest T. caespitosa var. caespitosa. Illustration of leaf sections indicates abaxial leaf surface has a margin of only 8 smooth cells (no papillose cells); this is followed by a stomata band with 11 rows of stomata, and partially papillose midrib 13 cells wide, lacking papillae mostly on the center 5 rows.

SE Manchuria: Maximowicz, yr 1860. Illustration shows abaxial leaf margin to lack papillae across 6 marginal cells followed by 8 papillose cells, a midrib of 20 cells wide, densely papillose, and stomata in 10 rows. The thick branchlets and phyllotaxy compare favorably with the preceding specimens, but as noted these show similarities to T. baccata. Here the leaf section characteristics also compare more closely to T. baccata. The geographical location is more northern and western. Additional data are needed to support any determination that this and the other specimens of this intermediate group should be classified under T. baccata. Other specimens from Maximowicz reportedly from this region include T. umbraculifera var. umbraculifera and T. umbraculifera var. microcarpa.

Cultivar: Secrest Arboretum, Wooster, OH, appearing to be a hybrid, T. caespitosa var. latifolia x T. umbraculifera