Pinatae .
Pinaceae 319
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© Springer-Verlag Berlin Heidelberg 1990
Pinaceae
C. N. PAGE
Pinaceae Lindl., Nat. Syst. Bot. ed. 2: 313 (1836).
Highly resinous, monoecious shrubs or trees. Leaves
deciduous or evergreen, usually with resin canals, aci-
cular, linear or oblong, solitary and spirally arranged
on all shoots, or in some genera only so on new
growth, and later densely grouped on short, lateral,
spur shoots, soft, stiff, flexible, rigid or asperous in tex-
ture. Shoots glabrous, pubescent or hairy, usually with
distinctive, rhythmic annual growth increments, and
scaly, terminal, often resinous buds in the resting sea-
son. Male and female cones usually on the same tree,
the female often appearing later in the life of the tree
or borne mostly on higher branches. Male (pollen)
cones catkin-like, solitary or grouped, axillary, each
cylindric with numerous, spirally-arranged, microspor-
ophylls with two pollen sacs on the lower surface of
each. Female (ovuliferous) cones solitary, in irregular
groups or whorled, erect, suberect, or pendant, sessile
or shortly-stalked, rounded or elongate, frequently
large, each with numerous spirally-arranged scales and
more or less conspicuous bracts, the former each with
two ovules on their upper (adaxial) surface, leathery or
fleshy at first, becoming stiff, woody, flexible, papery
or leathery with age, and sometimes highly resinous,
maturing in 1- 3 seasons and eventually dismembering
on the tree or the scales opening on the tree to release
the seed. Seed winged or unwinged, two to each scale.
Cotyledons of variable number, sometimes numerous.
A family of probably 12 genera and about 200 species,
almost exclusively in the northern hemisphere.
VEGETATIVE ANATOMY AND MORPHOLOGY. Plants of
this family are mostly fast-growing shrubs or usually
trees of moderate to large size, sometimes attaining a
very considerable age. Most are of more or less mono-
podial habit when young, but this habit becomes more
irregular with age in most genera, other than Larix. Pi-
cea. and Abies. Particularly broadly-crowned ultimate
trees can occur in Cedrus and some Pinus (e. g., P. at-
tenuata. P. pinea) and Keteleeria. whilst in Pseudolarix.
Cathaya. Pseudotsuga. Hesperopeuce, and Tsuga, the
crown shape, very strongly influenced by light factors
and the density of crowding of populations, may also
be broad. The trunk is usually simple, erect and
columnar, sometimes slightly buttressed, with thin or
moderately thick bark that eventually falls in irregular
plates and scales. Bark characters vary from very
rough and thick (especially in Pseudotsuga and some
Pinus) to fairly smooth (in most Abies). The branches
are whorled, at least when young or throughout much
320 Pinatae . Pinaceae
of the life of the tree, and persistent or shed from the
trunk with age.
Seedlings of all genera have free, simple, spreading,
flattened leaves arranged radially around the shoot.
Where the adult foliage is different from this, the
differences usually appear rapidly once the plants
progress beyond the immediate seedling stage.
Leaves are simple in all genera. In Pinus they are
fasic1ed. They are notably dorsiventrally flattened in
Pseudolarix, Larix, Cathaya, Pseudotsuga, Keteleeria,
Tsuga, some Picea (e. g., P. brachytyla, P. omorika) and
most Abies. Leaves of various other cross-sectional
shapes occur in Cedrus, Hesperopeuce, several Picea
and some Abies, and Pinus.
Leaves are winter-deciduous in Pseudolarix and Larix.
Rather prominent leaf cushions occur in Cathaya, No-
thotsuga, Tsuga, Hesperopeuce, and Picea.
Short, lateral spur shoots are a distinctive feature of a
few genera (Pseudolarix, Cedrus, Larix), and Cathaya
and Hesperopeuce are progressively more subdimor-
phic in this respect. In all genera, growth ceases for a
period (normally the winter period) in the year, and
resting buds are laid down annually in both evergreen
and deciduous genera. The winter buds are markedly
elongate in Pseudotsuga.
Species with leaves which are flattened are more or
less strongly hypostomatic; those with leaves of other
shapes are usually amphistomatic. Stomata are fre-
quently in multiple linear rows, and where leaves are
hypostomatic, the rows are frequently set within shal-
low grooves either side of the leaf midrib, often
marked by encrustations of white wax.
One genus, Pinus, contains the world's oldest living
vascular plant individual: Pinus longaeva D. K. Bailey
(Bailey 1970).
For other details, see family and generic descrip-
tions, the following sections, and Bailey (1909), Hill
and de Fraine (1909), Jeffrey (1905, 1925), Morikawa
and Morikawa (1929), Harlow (1931), Lacassagne
(1934), Gaussen (1966), Maheshwari and Biswas
(1970), and Jain (1976).
POLLEN MORPHOLOGY. In the Pinaceae, there are three
pollen types: (1) the bisaccate Pinus type, (2) the Tsuga
type, restricted to this genus, in which the pollen is
provided with a circular, inflated frill and a distinctive
SCUlpturing at the proximal pole, and (3), the Larix
type, characterized by a smooth pollen surface and the
absence of sacci and frills, and very large grains, re-
stricted to Larix and Pseudotsuga.
KARYOLOGY. The chromosome base number of n= 12
appears to be widespread in most genera of the Pina-
ceae. This base number distinguishes this family from
n= 11 in both the Taxodiaceae and Cupressaceae. The
only wholly anomalous pinaceous genus in base num-
ber appears to be Pseudolarix, with n=22, 2n=44.
REPRODUCTIVE BIOLOGY. All genera are wind-pollinat-
ed. Pollination takes place in early spring in most gen-
era. Seed matures in the same season, or in one or
more subsequent seasons in Pinus.
Male and female cones appear on the same tree.
Most genera reach male maturity in the life of each
tree simultaneously with, or earlier than female matu-
rity. Female cones occur widely scattered in the trees
amongst males in Pseudolarix, Cedrus, Larix, Cathaya,
Hesperopeuce, Pseudotsuga, Tsuga, and many Pinus, or
are grouped more usually on the uppermost branches
of the tree in Keteleeria, Picea, Abies, and some Pinus.
Differences between maximum pollen shedding
times and female receptivity probably occur in most
species, promoting outbreeding. In those genera in
which pollen is winged, the air-bladders probably as-
sist pollen orientation on the micropyle.
Mature female cones are notably erect in Cedrus,
Keteleeria, Abies, and perhaps Nothotsuga, and sub-
erect, variously orientated, or pendulous in Pseudola-
rix, Larix, Cathaya, Pseudotsuga, Nothotsuga, Tsuga,
Hesperopeuce, Picea, and Pinus. In Picea they are erect
at fertilization, and invert subsequently; and a similar
inversion, developed to a lesser degree, characterizes
also Hesperopeuce.
Seed is shed gradually either by the parting of cone
scales at maturity in Larix, Cathaya, Pseudotsuga, Ke-
teleeria, Nothotsuga, Hesperopeuce, Tsuga, Picea, and
Pinus, or by dismemberment of the whole cone in
Pseudolarix, Cedrus, and Abies. In Cedrus and Abies,
the central columella of the cone is usually left at-
tached to the shoot. In Keteleeria seeds may be long-
retained. In Pinus, cones of several species are seroti-
nous, and open only following stimulation by fire.
Pinus contains the largest female cones and the
heaviest in the family. Seeds are winged throughout
most of the family, except in a few species of Pinus.
SUBDIVISIONS AND RELATIONSHIPS WITHIN THE FAMILY.
Although the family Pinaceae has been traditionally
subdivided, there is lack of general agreement between
authors on how this should be done. Perhaps the most
distinctively separable group are the true pines (Pinus),
which have often been removed as a separate subfami-
ly from the remainder of the family (see comments un-
der "Interrelationships between families").
The pines apart, this situation reflects the diversity
of interrelationships which appear to exist between
many of the genera. Although this topic, as in almost
all other conifer families, is much in need of further
study, accumulating evidence suggests that the pat-
 Pinatae . Pinaceae
terns of intergeneric relationships are extremely com-
plex and diverse, perhaps involving intergeneric hybri-
dization as well as evolutionary radiation and the
survival of archaeic and sometimes generically inter-
mediate taxa. Indeed, the diversity of intergeneric
links that appear to exist between many genera of the
Pinaceae lend little support to the necessity of creating
strict supergene ric subdivisions within the non-pine
parts of the family, as these, on biological and evolu-
tionary grounds, would seem to have a strong element
of artificiality.
Apart from Pinus, the genera Pseudolarix, Cedrus,
and perhaps Larix appear to stand as the most, al-
though not wholly, isolated within the Pinaceae. Picea
appears to show close links with Tsuga in both vegeta-
tive and reproductive features, and through it, per-
haps, to Pseudotsuga. Pseudotsuga may also link
distantly to Keteleeria and thereby to Abies in one di-
rection, especially in vegetative morphology, and to
Tsuga and Cathaya in another. Keteleeria also appears
to link to Pseudolarix in some aspect of its cone mor-
phology.
There has, however, been a lack of general agree-
ment in relation to the affinities and homogeneity of
Tsuga and its near allies, including Cathaya, Nothotsu-
ga, and Hesperopeuce. One view (e. g., Silba 1984,
1986) is to group all these genera into Tsuga. The fit of
several of them into this genus seems, however, un-
comfortable, and to group them there seems to over-
simplify the probable biological reality and possible
past history of their, apparently complex, generic in-
terrelationships.
The alternative view, adopted here, is to recognize
Cathaya, Nothotsuga. and Hesperopeuce as separate
genera which, whilst having clear affinities with Tsuga,
also have each of them, strong and distinctive affinities
with other pinaceous genera, which, for each, are not
the same, nor are they the same as those of the remain-
ing species of Tsuga. Thus removed, each genus, as
well as all remaining Tsuga, appear to become much
more wholly natural genera. Cathaya differs from Tsu-
ga especially in its subdimorphic to dimorphic shoot
structure, with much of its foliage clustered and more
or less whorled on short, lateral, spur shoots, and in
the possession of two-bladdered pollen grains, lacking
the characteristic frill found throughout the species of
Tsuga. Nothotsuga differs from Tsuga especially in its
long, acute-tipped winter buds, its long, acute leaves
which, although flattened, are hypostomatic, and pe-
dunculate female cones with long, exerted, subspathu-
late bracts. The female cones are also reported to be
erect. Hesperopeuce differs from Tsuga especially in its
thick, angular, assurgent, amphistomatic leaves, its
subdimorphic shoot morphology, and its long, ovoid-
cylindric cones with reflexing basal scales, the cones
being suberect before and long after pollination, and
321
the dark, furrowed bark of its trunk. Hesperopeuce also
has distinctly bladdered pollen grains, a feature not
found in Tsuga. Possible links between these genera
and others seem diverse. Gaussen (1966) points to
some intermediacy of Cathaya between Tsuga and Pi-
nus, and it has also been suggested to resemble Pseu-
dolarix in its shoot morphology, Keteleeria in its leaf
morphology, and Pseudotsuga in its female cone. The
bladdered pollen grain of Cathaya is of the Pinus type,
and this feature, plus its subdimorphic shoot morphol-
ogy, links Cathaya and Hesperopeuce, which in many
respects seem to be the trans-Pacific equivalents of one
another. Gaussen (1966) viewed Hesperopeuce as so in-
termediate between Tsuga and Picea as to suggest it to
be a recent hybrid. Although affinities do exist with
both these genera, and especially with Picea in its fe-
male cone, the plant seems more likely a palaeo-relict,
and in many aspects, notably its shoot morphology,
leaf structure and arrangement, mature tree habit, and
bark characters, Hesperopeuce contains conspicuous
affinities with Cedrus, a link not hitherto suggested.
Nothotsuga, with its distinctive and possibly erect,
bracteate female cone seems also of highly diverse and
enigmatic affinity. Gaussen (1966) regarded it as inter-
mediate between Tsuga and Keteleeria, whilst it also
resembles Pseudotsuga in its exerted (though not trifid)
bracts. It seems possible, too, that its bracteate and
possibly erect female cone, might link it also with
more primitive members of Abies.
The very large and distinctive genus Pinus, probably
the evolutionarily most advanced in the family, stands
much by itself, and has been the topic of many studies
of its own (e. g., Mirov 1967). Its winged pollen is a
possible link with that of such genera as Cathaya,
whilst also in eastern Asia, Pinus krempfii [Ducampopi-
nus krempfii (Lecomte) A. Chev.] of Vietnam and Cam-
bodia, with highly flattened leaves, seems primitive
within the genus, and has been suggested to show dis-
tinct affinities perhaps with Pseudolarix or Keteleeria.
Aspects of species interrelationships within one of
the large genera (Picea) have been considered by
Wright (1955) and Page and Hollands (1987).
AFFINITIES OF THE PINACEAE. The Pinaceae are a dis-
tinctive family of conifers, without close alliances with
other living conifer families, and with a relatively long
fossil history.
Fossils clearly referable to the Pinaceae, including
those of pollen, ovulate cones, seeds, imprints of ve-
getative organs, and petrified wood, occur widely in
the northern hemisphere in sediments of Late Meso-
zoic and Tertiary age. This fossil record indicates that
many, if not all, of the modern genera of the Pinaceae
had evolved by the end of the Oligocene, with much of
their evolution probably taking place in the Late Meso-
zoic and Early Tertiary (Miller 1976). Subsequently,
322 Pinatae . Pinaceae
most suffered range restriction with the onset of the
Pleistocene.
All genera include living and fossil species which
are extensively or very largely northern hemisphere in
distribution, and probably almost all of the evolution
of genera and species has taken place in this hemi-
sphere.
DISTRIBUTION AND ECOLOGY. The Pinaceae remain an
almost exclusively northern family of conifers, with
the majority of the species of all the large genera (not-
ably Pinus, Picea, Abies, and Larix) widely distributed
throughout the temperate parts of both Old and New
Worlds and extending into high northerly latitudes.
There are particular concentrations of species of each
of these genera in both North America and in the east
of Asia, with a considerable number of endemic spe-
cies of more restricted range in the Sino-Himalayan
region. Tsuga and Pseudotsuga follow a similar pat-
tern, but are in North America and eastern Asia only,
and are absent from Eurasia. Hesperopeuce is re-
stricted to western North America. Cathaya, Nothotsu-
ga, and Keteleeria are restricted to the Sino-Himala-
yan region (the former two entirely to China), and Ce-
drus ranges discontinuously from the Mediterranean
basin to the western Himalayas. Apart from one spe-
cies of Cedrus in Algeria and Marocco, all species of
Pinaceae are absent from Africa and none reach South
America or Australasia. Most of the limited number of
species which spread southward in Central America or
in SE Asia, are essentially montane.
Most species of all genera in the family are trees of
generally poor, acidic and either wet or rocky habitats,
sometimes forming mixed evergreen or evergreen-
broadleaved forests, but more often forming extensive
monotypic stands over large, north-temperate areas.
Northern species of several genera (notably Abies and
Picea) adopt a particularly narrow, monopodial habit,
which sheds snow readily. The limited modern distri-
bution of one genus, Keteleeria, appears to be that of
regularly cyclone-damaged vegetation. A few species
of some genera (such as Picea glehnii) are specialists of
certain soils of ultrabasic character yielding heavy
metals. Some species of most genera grow on lime-
stone soils (notably so in China and upper Burma).
Many species of Pinus are tolerant of poor sandy soils.
A few, such as Pinus thunbergii, are also tolerant of di-
rect exposure to salt-laden air, and several (notably
those of the Mediterranean and California) are fire-
tolerant or require fire for seed release. Probably all
genera of Pinaceae in the New World and at least Pi-
nus, Abies, and Keteleeria in the Old World contain
some species (in North America, especially western
species) which are parasitized by dwarf mistletoes of
the genus Arceuthobium (Viscaceae), and may be
sometimes heavily infected (Hawkesworth and Wiens
1972).
Amongst other parasites, rust fungi of the order
Uredinales occur widely on the Pinaceae, the latter as
aecial hosts, and including the genera Pseudotsuga,
Larix, Pinus, Cedrus, Abies, Picea, and Tsuga (e. g.,
Durrieu 1980).
Other ecological, phytogeographic or floristic
aspects in regions where the Pinaceae are especially
abundant have been discussed by Wilson (1916), Sud-
worth (1916), and Kuan (1981), and aspects of their
fossil distribution especially by Dubois- Ladurantie
(1941), Ferguson (1967), and Miller (1976).
KEy TO GENERA OF PINACEAE.
1. Leaves deciduous 2
- Leaves evergreen 3
2. Mature female cones dismembering on tree 2. Pseudolarix
- Mature female cones remaining whole and falling intact
1. Larix
3. Leaves united in groups of usually 2, 3, or 5 by sheaths at
the base 12. Pinus
- Leaves not united in groups 4
4. Leaves distant on long shoots and more closely clustered
on short, side shoots 5
- Leaves all distant on long shoots; short, side shoots ab-
sent 7
5. Mature female cones dismembering on tree 10. Cedrus
- Mature female cones remaining whole and falling intact 6
6. Leaves flattened, hypostomatic 6. Cathaya
- Leaves not flattened, amphistomatic 8. Hesperopeuce
7. Shoots roughened after leaf abscission by persistent leaf
bases 8
- Shoots ± smooth, leaf bases not persistent 10
8. Leaves sessile, mostly pointed and usually pungent
11. Picea
- Leaves shortly and distinctly petiolate, never pungent 9
9. Bracts of female cones small, not exerted 9. Tsuga
- Bracts of female cones large, exerted 7. Nothotsuga
10. Winter buds spindle-shaped, acute; female cones pendu-
lous or variously orientated 5. Pseudotsuga
- Winter buds ovoid, blunt; female cones erect 11
11. Mature female cones dismembering on tree 4. Abies
- Mature female cones remaining whole and falling intact
3. Keteleeria
1. Larix Mill. Fig. 167
Larix Mill., Gard. Diet. Abr. ed.: 4 (1754).
Tall, monoecious, deciduous trees. Leaves linear, re-
mote, and spirally arranged on long, leading shoots,
densely clustered and whorled on short, lateral spur
shoots. Male cones solitary, frail, ovoid-cylindric. Pol-
len not winged, smooth. Female cones solitary, subglo-
bose to ovoid-oblong, with long, simple bracts which
are hidden or protruding at maturity.
About ten species widely distributed around the north-
temperate zone.
 Larix· Pseudolarix· Keteleeria . Abies
2. Pseudo/am Gordon Fig. 168
Pseudolarix Gordon, Pinet.: 292 (1858).
Chrysolarix H. E. Moore, Baileya 13: 133 (1965).
Tall, monoecious, deciduous trees. Leaves linear,
pointed, remote and spirally arranged on leading
shoots, whorled on ± club-shaped, lateral spur shoots.
Male cones in umbellate clusters, terminal on short,
lateral, spur shoots. Pollen with 2 air-bladders. Female
cones ovoid-pyramidal, brittle and deciduous at matu-
rity.
A single species of the mountains of Central and NE
China, in Zheziang, Jiangsu, Guangxi, and Anhui pro-
vinces.
3. Keteleeria Carriere Fig. 169
Keteleeria Carriere, Rev. Hart. 1866: 449 (1866); Farjon,
Notes Roy. Bot. Gard. Edinburgh 46: 81-99 (1989).
Tall, monoecious evergreen trees. Leaves linear, flat-
tened, spirally arranged, mostly spreading into 2, ±
strongly pectinate ranks, eventually leaving ± smooth-
er, circular leaf scars. Male cones in terminal or axil-
lary umbels. Pollen with 2 air-bladders. Female cones
strongly erect, elongate-cylindric; bracts small, and in-
conspicuous, not protruding beyond the scales at mat-
urity.
323
c D
Fig. 167 A-G. Pinaceae. Larix decidua. A Long shoot with
spur shoots. B Branch with buds. C Ovuliferous scale.
D Male cone. E Seed cone. F Bract and seed scale from be-
low. G Seed scale with seeds (Original G6tz)
A genus of three to seven somewhat ill-defined spe-
cies, endemic to warm-temperate regions of China,
Hong Kong, Taiwan, Laos, and Vietnam; widespread
in China but scarce and local elsewhere; known in Eu-
rope as fossils of Tertiary age.
4. Abies Mill. Fig. 170
Abies Mill., Gard. Diet. Abr. ed.: 4 (1754); Farjon and Rush-
forth, Notes Roy. Bot. Gard. Edinburgh 46: 59-79 (1989).
Tall, monoecious, evergreen trees. Leaves linear, most-
ly ± flattened, rarely 4-angled, spreading all around
the shoot or twisted at the base and variously arranged
into lateral ranks, leaving ± smooth, circular, leaf
scars on the shoot. Male cones grouped, axillary. Pol-
len with 2 air-bladders. Female cones strongly erect,
elongate-cylindric, the bracts protruding or hidden at
maturity, the scales and bracts eventually dismember-
ing and falling from the central cone axis which re-
mains long-persistent on the branches.
A large genus of over 40 species, widely distributed
through the north-temperate zone. Revision and subdi-
vision: Liu (1971); Viguie and Gaussen (1929).
324 Pinatae . Pinaceae

Fig. 168A-F. Pinaceae. Pseudolarix amabilis. A Branch with

seed cone. B Branch with male cones. C Ovuliferous scale,
seen from below. D The same, seen from above. E Male
cone. F Seed (A from Fitschen 1930; B from Rendle 1971;
others from Koehne 1893)
 Pseudolarix· Keteleeria . Abies 325

(:;)
Dr
I nI
I I
iI \ I' II
.&
Fig. 169 A-F. Pinaceae. A, B Keteleeria fortunei. A Sterile . IiiI

branch. B Branch with male cones. C-F Keteleeria davidia-

I\~
na. C Female cone. D Seed scale with bract from below.
E The same from above. F Seed (Original Gotz)

J .
v
Fig. 170A-J. Pinaceae. A Abies grandis, underside of~
branch. BAbies koreana , leaf, transversal section, upper and
lower side. C-E Abies alba . C Male cone. D Microsporo-
phyll. E Bract and scale. F-I Abies koreana. F Seed cone,
G Scale with seed, upper view. H $cale with bract, lower
view. I Seed. J Abies venusta, bract and scale (all original
Gotz)
326 Pinatae . Pinaceae
c D
B
Fig. 171 A-G. Pinaceae. Pseudotsuga menziesii. A Branch.
8 Male strobilus. C Microsporophyll. D Female cone.
E Bract and seed scale. F Seed scale with seeds. G Seed
(A Original Giitz ; 8 from Flora Iberica 1986; C from
Fitschen 1930)
5. Pseudotsuga Carriere Fig. 171
Pseudotsuga Carriere, Tr. Gen. Conif. ed. 2: 256 (1867).
Tall, monoecious, evergreen trees. Leaves linear, flat-
tened, spirally arranged, spreading or in 2, ± subpec-
tinate ranks, eventually leaving very slightly raised,
transversely oval, leaf scars. Winter buds elongate,
acute, lustrous, non-resinous. Male cones solitary, axil-
lary. Pollen not winged, smooth. Female cones solitary
or grouped, ovoid-oblong, with long, acute-tipped
bracts protruding beyond the scales.
At least four to seven species distributed around the
north Pacific, in Japan, China, Taiwan, and western
North America south to Mexico. Those of China are
somewhat ill-defined and poorly known.
G
6. Cathaya Chun & Kuang Fig. 172
Cathaya Chun & Kuang, Bot. Zh. SSSR. 43: 464 (1958).
Tall, monoecious, evergreen trees. Leaves linear, flat-
tened, abundant and spirally arranged, borne from
slightly prominent leaf cushions on long leading
shoots; clustered and ± whorled on short, lateral spur
shoots. Male cones solitary, terminal on short, lateral
spur shoots. Pollen with two air-bladders. Female
cones solitary, broadly ovoid-oblong; the bracts not
protruding beyond the scales at maturity.
An obscure genus of one or possibly more rather little
known, relict living species, endemic to western China
(Guangxi, Hunan?, and Sichuan), with fossil (Tertiary)
representatives also elsewhere. Although its generic
status has been sometimes doubted, Cathaya seems to
link variously with Pseudolarix, Tsuga, Pseudotsuga,
and perhaps Keteleeria.
7. Nothotsuga H.-H. Hu & c. N. Page
Nothotsuga H.-H. Hu & c. N. Page, Notes R. Bot. Gard.
Edinburgh 45: 390 (1989).
Tall, monoecious, evergreen trees. Leaves narrowly lin-
ear-elliptic, long, flattened, spreading into 2, somewhat
 Hesperopeuce· Tsuga . Cedrus
contorted ranks, set on shining projections with promi-
nent cushion-like bases. Male cones clustered, termi-
nal. Female cones solitary, oblong-ovoid, erect, the
bracts exerted beyond the scales at maturity.
A single species, Nothotsuga longibracteata (Cheng)
H.-H. Hu ex C. N. Page, endemic to Hunan, Guizhou,
Guangdong, and Guangxi. Although its generic status
has been sometimes doubted, Nothotsuga seems to
have affinities with both Tsuga and Keteleeria, as well
as perhaps with more primitive members of Abies.
8. Hesperopeuce(Engelm .) Lemmon Fig. 172
Hesperopeuce (Engelm.) Lemmon, Bienn. Rep. Cal. State
Board For. 3: 126 (1890).
Tall, monoecious, evergreen trees. Leaves long, curved
and thick, ± bevelled at the tip, radially arranged and
assurgent, with cushion-like bases (pulvini) persistent
after the leaves fall. Male cones solitary, axillary in a
few, widely scattered leaves of the previous season.
Pollen with 2 air-bladders. Female cones solitary, ses-
sile, ovoid-cylindric, the basal scales often becoming
strongly recurved at maturity, the bracts not protrud-
ing beyond the scales at maturity.
A single species, Hesperopeuce mertensiana (Bongn.)
Rydb. (Tsuga mertensiana (Bongn.) Carriere), endemic
to western North America from southern Alaska to
central California, especially at the snowline. Al-
though its generic status has been sometimes doubted,
Hesperopeuce seems to link Tsuga, Picea, and Cedrus.
327
Fig. 172A-E. Pinaceae. A-D Calhaya argyrophylla.
A Branch with female cones. B Seed scale, lower view, with
bract. C The same, upper view, with seeds. D Seeds. E Hes-
peropeuce mertensiana. branch with female cones (A-D from
lconogr. Corm. Sin. 1972; E from Fitschen 1930)
9. Tsuga Carriere Fig. 173
Tsuga Carriere, Tr. Gen. Conif.: 83 (1847).
Tall, monoecious, evergreen trees. Leaves short, linear,
straight, the apex often notched, dorsiventrally flat-
tened, the apex spreading into 2, ± strongly pectinate
ranks, set on shining projections with cushion-like
bases (pulvini), persistent after the leaves fall. Male
cones subglobular, solitary, axillary. Pollen not winged,
but with an annular frill. Female cones terminal on lat-
eral shoots of previous year, solitary, broadly ovoid,
the bracts small, and inconspicuous.
About 14 species in eastern and western North Ameri-
ca and the far east of Asia (China, Japan, Taiwan).
10. Cedrus Trew Fig. 174
Cedrus Trew, Cedro Libani Hist. (1): 6 (1757) (nom . cons.);
Link, Linnaea 15: 537 (1841).
Tall, monoecious, evergreen trees. Leaves needle-like,
spirally arranged on long, leading shoots, whorled on
lateral spur shoots. Male cones solitary, robust, erect,
ovoid-cylindric, mostly abscissing from the shoots af-
328 Pinatae . Pinaceae

~
/.O;

(. .
.;0 H

Fig. 173A-H. Pinaceae. Tsuga canadensis. A Branch with fe-

male cone. B Bud. C Leaves, lower and upper side. D Ma-
ture female cone. E Seed scale with seeds. F Bract and seed
scale, lower view. G Male cone. H Microsporophyl\ (G, H
A from Koehne 1893; others original G6tz)

Fig. 174A-F. Pinaceae. Cedrus atlantica. A Long shoot with

spur shoots. B Tip of branch with buds. C Male cone.
o Branch with ovuliferous cone. E Seed scale and seeds.
F Seed (C-F from Flora Iberica 1986; others original G6tz)
 Picea . Pinus
ter maturity. Pollen with 2 air-bladders. Female cones
usually solitary, erect, ovoid-oblong, with a depressed,
obtuse apex.
Four species with disjunct distributions, on rocky
mountainsides from N. Africa (Atlas Mountains) to
the western Himalayas.
11. Picea A. Dietr. Fig. 175
Picea A. Dietr., Fl. Geg. Berlin 2: 794 (1824).
Tall, monoecious, evergreen trees. Leaves linear, acute,
needle-like, quadrangular or flattened, ± spreading
all around the shoot or twisted at the base and var-
iously arranged into lateral, usually regularly pectinate
ranks, set on woody, peg-like projections on cushion-
like bases (pulvini), persistent after the leaves fall.
Male cones grouped, axillary. Pollen with 2 air-blad-
ders. Female cones erect at first, pendulous at maturi-
ty, ovate-elongate, the bracts not protruding.
329
Fig. 175 A-J. Pinaceae. A, B Picea orientalis. A Branch.
B Branch, leaves removed. C-G Picea abies. C Male strobi-
lus. D Microsporophyll, lower view. E The same, lateral
view. F Bract and seed scale. G Ovuliferous scale. H-J Pi-
cea breweriana. H Seed cone. I Bract and seed scale. J Seed
scale with seeds (C, D from Fitschen 1930; E, F, G from
Wettstein 1935; others original Gotz)
About 40 species in the northern hemisphere, particu-
larly widely distributed in the cool temperate latitudes.
For a subdivision, see Lacassagne (1934) and Liu
(1982).
12. Pinus L. Figs. 176, 177
Pinus L., Sp. PI.: 1000 (1753).
Tall, monoecious, evergreen trees. Leaves vary from
small and scale-like to long and linear-Ianceolate to
very long and narrowly linear, variously distributed,
330 Pinatae . Pinaceae
E F G
Fig.176A-I. Pinaceae. A, B Pinus sylvestris. A Branch with
mature seed cone. B Spur shoot. C, D Pinus nigra. C Scale
with bract. D Scale with ovules. E-G Pinus sylvestris.
E Seed. F Scale with seeds. G Scale, lower view. H Pinus
cembra, seed cone. I Pinus strobus, seed cones (A from Le
Maout et al. 1876; B from Troll 1954; C, D from Wettstein
1924; H from Fitschen 1930; I from Herter 1939; others orig-
inal Gi:itz)
the leaves solitary and spirally arranged, or (mostly) in
fascicles of 1-5, with persistent or deciduous scarious
basal sheaths. Male cones each in the axil of a scale
leaf, numerous and closely clustered around the bases
of young shoots, with numerous scales. Pollen with
2 air-bladders. Female cones terminal or subterminal,
elongate to ovate, symmetric or oblique, mostly
woody.
At least 80 species, widely distributed in the northern
hemisphere, south to Central America, North Africa,
 Selected Bibliography
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co, Baja California, 1800 m. Phot. K. Kubitzki
the Philippines and Sumatra. Revision: Mirov (1967);
Bailey (1970) (regional).
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