18.
Taxus fastigiata Lindley, Syn. Brit. Flora
241 (1829). T. baccata
(var.) fastigiata (Lindley) Loudon, Arb. frutic. britt. 4: 2066
(1838). T. baccata f. fastigiata (Lindley) Pilger,
Planzenreich 18(iv, 5): 115. 1903. Taxus baccata var. hibernica Hook.
ex Henkel & Hochstetter, Syn. Nadelhölzer 356 (1865), no specimens
cited, none reported in the Lindley Herbarium. Type: Ireland,
“mountains of Fermanagh” (transplanted to Florence Court in 1780, or
1760?), Willis, specimen unknown. Neotype (designated, Spjut
2007b)—from Florence Court—ex Herb. Jackson, Stewart, Sep. 1890
( K!).
Taxus baccata (var.) columnaris Carrière, Traité gén.
conif. 738. 1867, taxonomic synonym proposed. Taxus baccata f. columnaris (Carrière )
Beissner, Handb. Nandelholzk. 170. 1891.
Original material unknown, reported from horticulture in
Versailles, France. Distinguished
by Carrière for its columnar habit with variegated color, and short
branchlets on leader shoots. Type
undetermined.
Taxus baccata
(var.)
compressa Carrière, Traité gén. conif. 738. 1867, taxonomic
synonym proposed. Taxus
baccata (f.) compressa (Carrière ) Beissner, Handb.
Nandelholzk. 171. 1891. Compared by Carrière to his var. intermedia and to
var. fastigiata. Described
by Ouden & Boom (1965) as dwarf, compact and conical in habit with
numerous branchlets. Type
undetermined.
Taxus baccata stricta
Lawson,
Agric. Man. 398 1836 (n.v.), taxonomic synonym proposed. Taxus baccata f. stricta
Rehder, Bibl. Trees 2. 1949. Treated
by Rehder as a synonym of the under the Irish yew.
Type unknown.
Irish
yew. Distribution: British Isles (Ireland, England?).
18a. var. fastigiata. Tree, columnar and tapering to
apex, or cylindrical in outline; branches and branchlets mostly ascending to erect; bud-scales
loosely attached, 3–4 seriate on young growth, soon deciduous,
chartaceous, pale brown to chocolate brown, occasionally basal scales
deltoid smooth or with distinct midnerve on upper third, 1–3 mm long.
Leaves lacking on lower parts of branchlets, crowded and overlapping
near apex of branchlets, spreading in a radial manner, recurved, usually
overlapping the branchlet near base of blade when pressed, broad linear,
straight or slightly falcate, dark green, 1.5–3.0 cm long, 2.0–3.0
mm wide, 200–350 µm thick, slightly convex above to a low rounded
midrib, plane to slightly concave below to a flush or low rounded
midrib, plane to slightly revolute along margin, truncate (boat-shaped)
in x-sect.; upper (adaxial) epidermal cells elliptical in transverse
section, 8–15 µm tall, 10 µm wide, thin-walled; lower (abaxial)
nonstomata epidermal cells similar in transverse section, 10–18 cells
across the marginal zone, slightly larger on midrib, nearly quadrate to
± short rectangular, or trapezoidal, 1.5–4.5× l/w, narrowest in 4–11
rows near margins, slightly inflated, papillose on one-third to nearly
halfway between margin and stomata band, or not at all papillose on
marginal cells, epapillose on midrib, or
obscurely papillose on outer rows of midrib cells; papillae positioned
medially on midrib and marginal cells, submarginal on accessory cells,
low and indistinct, ±concrescent in 1–2 alternate rows on each cell;
stomata bands broader than the marginal region, with 8–10 (-11) rows of stomata,
greenish. Cones not studied in detail.
The Irish yew is thought to be native
to mountains in Fermanagh
County (Cuilcagh Mountains) in northwestern Ireland
where two female plants were reportedly discovered by a farmer,
George Willis, in 1773, or earlier, who dug up the plants and planted
one in his garden and gave the other to his landlord, who subsequently
planted it at Florence Court (Veitch et al. 1881, with reference to The Gardeners Chronicle 1873: 1336
where further reference is made to
undated manuscript by Lord Kinnaird); the one at Florence Court, which
has been the source for "millions of yew plants distributed throughout
the world" (Veitch et al. 1881), still survives (Internet sources,
2006), whereas Willis' plant died in 1865 (Veitch et al. 1881). Additionally, Loudon
(1838 with reference to MacKay, Flora Hibernica: 260) had suggested that Irish yew was in cultivation at Comber in
the county of Down, and near the town of Antrim before 1780; an
illustration of one such plant in Loudon (1838, shown below) was
indicated to have been provided by MacKay, and which was accompanied by
a description from the proprietor of the tree ( C. J. Andrews), and
furthermore, Loudon (1838) implied that the
original tree still existed in the mountains near Florence Court.
For many years it was thought that there were only female plants of the
Irish yew (Loudon 1838), but male plants were later discovered at North Mundham in
Sussex, England (Bean 1953; see also letter from Fletcher below
accompanied by specimen collected in 1897). One might wonder
whether these male plants came from another
source, but it is generally known that yew plants can occasionally reverse
their sex, or can be monoecious to a limited extent as
seen in the Dovaston yew.
The Irish yew (T. fastigiata) has long been considered
distinct from other yews for its erect habit and dark green leaves in
whorl-like arrangement; yet, most authorities treat it as a
synonym of T. baccata. This is because plants
reportedly grown from seed of Irish yew, which are thought to have been
pollinated by "common yew," are often regarded as the same as the "common yew"
(Veitch et al. 1881)
However, the Irish yew is viewed by this author as having ancestral
features worthy of species status even though it may form hybrids. In many specimens of the Irish yew,
leaves appear distinct for their boat-like shape in cross section, especially the
sharply angled margins; no other
species of yew show this. Additionally, the abaxial epidermal cells often lack
papillae across a relatively broad marginal region, and often
entirely across the midrib as well, a feature that is generally seen in the
Cuspidata Alliance.
The radial orientation of leaves that
characterizes the Irish yew
is also much like the E Asian T. umbraculifera. They
differ in the manner in which leaves spread from branches.
In the Irish yew, the leaves spread by curving of the blade downwards
from base towards apex, whereas in the umbrella yew, the leaves bend more
abruptly in that they are sharply (reflexed) near the petiole. Other species,
which have radial orientation of leaves—the E Asian species, T.
caespitosa and the European T. baccata var. ericoides
(Morocco)—differ in the leaves curving upwards instead of downwards.
However, some specimens in the T. cuspidata
Alliance—from the islands of Sakhalin, Hokkaido, and Honshu—have been
difficult to distinguish from the Irish yew.
For example, I annotated a specimen collected by Dvorakovskia
& Bokina (A) from
Sakhalin Island as T. fastigiata
in June 1996, because of leaves noticeably recurved and
dark green in color similar to
that of the Irish yew. Branching, leaf
arrangement and leaf anatomy of this specimen also compared favorably with T. baccata var. glauca,
especially in view of another specimen
from Sakhalin Island, Flanakan &
Kirkham 203 (K), that appeared more similar to T. baccata
var. glauca. Specimens such
as these appear
infrequent, and while they have been difficult to resolve taxonomically, they have been recently
interpreted as belonging to T. caespitosa var. latifolia, which
is common on the islands of Hokkaido and Honshu.
Radial arranged
leaves, which are generally associated with plants having an erect
habit, are rare in Taxus. In the Euro-Mediterranean, most
specimens with these features
occur
near the western distribution limits of the genus. Taxus fastigiata,
for instance, is known primarily from two plants found in northwestern
Ireland.
Near
the southwestern range of the genus is Taxus baccata
var. ericoides, known only from middle Atlas of Morocco
(near Ifrane, 1700 m). Other
varieties of Taxus baccata, which also have limited geographical
occurrences, exhibit a south to north clinal relationship in
phyllotaxy with the leaf arrangement varying from radial to distichous.
Radial types include Taxus baccata
var. erecta found only in Spain and Portugal and
Taxus baccata var. pyramidalis occurring sporadically in Algeria,
Spain, France, and Norway.
The more common leaf arrangement—spreading horizontally in
one plane so as to appear in two-ranks—as exemplified by T. baccata var. baccata
and T. baccata var. washingtonii—may have evolved,
however, by introgression with ancestors related to T. biternata
and Taxus canadensis.
Introgression between Taxus
fastigiata and
T. recurvata is also evident with variation appearing
to correspond to a latitudinal cline instead of a south-north longitudinal
cline mentioned above. Plants of T. recurvata with more leafy
branches, and with leaves also showing more radial orientation, occur in the British
Isles (e.g. cv., 'Expansa'), in contrast to less leafy plants found
mostly in central and eastern Europe to southwest Asia
(e.g., T. recurvata var. linearis). Taxus
fastigiata var. sparsifolia, an endemic to the British Isles,
may also prove to be a hybrid between T. fastigiata and T.
recurvata; however, the introgression and the geographical
cline that is evident, may be a product of former contact with ancestral
Taxus contorta instead of T. recurvata. Many
specimens of Taxus recurvata from the Euro-Mediterranean were
found to have reddish parenchyma cells in the spongy mesophyll, a
feature that is more pronounced and characteristic of Taxus contorta.
Thus, while one may speculate that
Taxus contorta may have retreated into Europe since the Pliocene, it
may also have found refuge in the northwest Himalayas
as a result of the Himalayan uplift. Molecular studies, thus, may
help determine to what extent Taxus fastigiata, T. canadensis
and T. contorta have contributed to the taxonomic complexity of
yew in the Euro-Mediterranean.
The specimens shown below undoubtedly
include hybrids for which I do not make any further distinction;
however, the Hick's yew, which has a similar habit, and which may be
a hybrid involving T. fastigiata, is treated as a distinct variety
of Taxus umbraculifera based specimens that appear to have been
collected in the wild of Japan and are indistinguishable from its type.
Representative Specimens—Ireland: Florence Court, Stewart, Sep.
1890 (K); ex Herb. Lange (K). Cultivation—Europe—England:
Fletcher, 13 Mar. 1897 (K); Yorkshire with specimen ex Herb.
Baker, Oct. 1837 (BM). Portugal: Rainba 1873 (S: C-2069).
Switzerland: f. aurea, C. Baenitz s.n (US: 139642).
Pinetum Boissier, Genf, 1953, ex Herb. Dendrol. Schneider, Jeps. s.n.
(A). Granvik, Sep. 1886 (BM). Ex Herb. Zuccarini, annotated T.
hybernica (M). Boorman s.n. (A). North
America—U.S.A.: Oregon: near Salem, O Brien C, D, F, G
(wba)
|