central cylinder of the stem, and does not assume the girdle-like form
characteristic of the cycadean leaf-trace. In species of which the
leaves have more than one vein (e.g. Araucaria imbricata, &c.) the
leaf-trace leaves the stele of the stem as a single bundle which splits
up into several strands in its course through the cortex. In the wood
of some conifers, e.g. Araucaria, the leaf-traces persist for a considerable
time, perhaps indefinitely, and may be seen in tangential
sections of the wood of old stems. The leaf-trace in the Coniferales
is simple in its course through the stem, differing in this respect from
the double leaf-trace of Ginkgo. A detailed account of the anatomical
characters of conifers has been published by Professor
D. P. Penhallow of Montreal and Dr. Gothan of Berlin which
will be found useful for diagnostic purposes. The characters of
leaves most useful for diagnostic purposes are the position of the
stomata, the presence and arrangement of resin-canals, the structure
of the mesophyll and vascular bundles. The presence of hypodermal
fibres is another feature worthy of note, but the occurrence of these
elements is too closely connected with external conditions to be of
much systematic value. A pine needle grown in continuous light
differs from one grown under ordinary conditions in the absence of
hypodermal fibres, in the absence of the characteristic infoldings of
the mesophyll cell-walls, in the smaller size of the resin-canals, &c.
The endodermis in Pinus, Picea and many other genera is usually
a well-defined layer of cells enclosing the vascular bundles, and
separated from them by a tissue consisting in part of ordinary parenchyma
and to some extent of isodiametric tracheids; but this
tissue, usually spoken of as the pericycle, is in direct continuity with
other stem-tissues as well as the pericycle. The occurrence of short
tracheids in close proximity to the veins is a characteristic of coniferous
leaves; these elements assume two distinct forms—(1) the short
isodiametric tracheids (transfusion-tracheids) closely associated with
the veins; (2) longer tracheids extending across the mesophyll at
right angles to the veins, and no doubt functioning as representatives
of lateral veins. It has been suggested that transfusion-tracheids
represent, in part at least, the centripetal xylem, which forms a
distinctive feature of cycadean leaf-bundles; these short tracheids
form conspicuous groups laterally attached to the veins in Cunninghamia,
abundantly represented in a similar position in the leaves of
Sequoia, and scattered through the so-called pericycle in Pinus,
Picea, &c. It is of interest to note the occurrence of precisely similar
elements in the mesophyll of Lepidodendron leaves. An anatomical
peculiarity in the veins of Pinus and several other genera is the continuity
of the medullary rays, which extend as continuous plates from
one end of the leaf to the other. The mesophyll of Pinus and Cedrus
is characterized by its homogeneous character and by the presence
of infoldings of the cell-walls. In many leaves, e.g. Abies, Tsuga,
Larix, &c., the mesophyll is heterogeneous, consisting of palisade and
spongy parenchyma. In the leaves of Araucaria imbricata, in which
palisade-tissue occurs in both the upper and lower part of the
mesophyll, the resin-canals are placed between the veins; in some
species of Podocarpus (sect. Nageia) a canal occurs below each vein;
in Tsuga, Torreya, Cephalotaxus, Sequoia, &c., a single canal occurs
below the midrib; in Larix, Abies, &c., two canals run through the
leaf parallel to the margins. The stomata are frequently arranged in
rows, their position being marked by two white bands of wax on the
leaf-surface.
The chief home of the Coniferales is in the northern hemisphere, where certain species occasionally extend into the Arctic circle and penetrate beyond the northern limit of dicotyledonous trees. Wide areas are often exclusively occupied by Distribution. conifers, which give the landscape a sombre aspect, suggesting a comparison with the forest vegetation of the Coal period. South of the tree-limit a belt of conifers stretches across north Europe, Siberia and Canada. In northern Europe this belt is characterized by such species as Picea excelsa (spruce), which extends south to the mountains of the Mediterranean region; Pinus sylvestris (Scottish fir), reaching from the far north to western Spain, Persia and Asia Minor; Juniperus communis, &c. In north Siberia Pinus Cembra (Cembra or Arolla Pine) has a wide range; also Abies sibirica (Siberian silver fir), Larix sibirica and Juniperus Sabina (savin). In the North American area Picea alba, P. nigra, Larix americana, Abies balsamea (balsam fir), Tsuga canadensis (hemlock spruce), Pinus Strobus (Weymouth pine), Thuja occidentalis (white cedar), Taxus canadensis are characteristic species. In the Mediterranean region occur Cupressus sempervirens, Pinus Pinea (stone pine), species of juniper, Cedrus atlantica, C. Libani, Callitris quadrivalvis, Pinus montana, &c. Several conifers of economic importance are abundant on the Atlantic side of North America—Juniperus virginiana (red cedar, used in the manufacture of lead pencils, and extending as far south as Florida), Taxodium distichum (swamp cypress), Pinus rigida (pitch pine), P. mitis (yellow pine), P. taeda, P. palustris, &c. On the west side of the American continent conifers play a still more striking rôle; among them are Chamaecyparis nutkaensis, Picea sitchensis, Libocedrus decurrens, Pseudotsuga Douglasii (Douglas fir), Sequoia sempervirens, S. gigantea (the only two surviving species of this generic type are now confined to a few localities in California, but were formerly widely spread in Europe and elsewhere), Pinus Coulteri, P. Lambertiana, &c. Farther south, a few representatives of such genera as Abies, Cupressus, Pinus and juniper are found in the Mexican Highlands, tropical America and the West Indies. In the far East conifers are richly represented; among them occur Pinus densiflora, Cryptomeria japonica, Cephalotaxus, species of Abies, Larix, Thujopsis, Sciadopitys verticillata, Pseudolarix Kaempferi, &c. In the Himalaya occur Cedrus deodara, Taxus, species of Cupressus, Pinus excelsa, Abies Webbiana, &c. The continent of Africa is singularly poor in conifers. Cedrus atlantica, a variety of Abies Pinsapo, Juniperus thurifera, Callitris quadrivalvis, occur in the north-west region, which may be regarded as the southern limit of the Mediterranean region. The greater part of Africa north of the equator is without any representatives of the conifers; Juniperus procera flourishes in Somaliland and on the mountains of Abyssinia; a species of Podocarpus occurs on the Cameroon mountains, and P. milanjiana is widely distributed in east tropical Africa. Widdringtonia Whytei, a species closely allied to W. juniperoides of the Cedarberg mountains of Cape Colony, is recorded from Nyassaland and from N.E. Rhodesia; while a third species, W. cupressoides, occurs in Cape Colony. Podocarpus elongata and P. Thunbergii (yellow wood) form the principal timber trees in the belt of forest which stretches from the coast mountains of Cape Colony to the north-east of the Transvaal. Libocedrus tetragona, Fitzroya patagonica, Araucaria brasiliensis, A. imbricata, Saxegothaea and others are met with in the Andes and other regions in South America. Athrotaxis and Microcachrys are characteristic Australian types. Phyllocladus occurs also in New Zealand, and species of Dacrydium, Araucaria, Agathis and Podocarpus are represented in Australia, New Zealand and the Malay regions.
Gnetales.—These are trees or shrubs with simple leaves. The flowers are dioecious, rarely monoecious, provided with one or two perianths. The wood is characterized by the presence of vessels in addition to tracheids. There are no resin-canals. The three existing genera, usually spoken of as members of the Gnetales, differ from one another more than is consistent with their inclusion in a single family; we may therefore better express their diverse characters by regarding them as types of three separate families—(1) Ephedroideae, genus Ephedra; (2) Welwitschioideae, genus Welwitschia; (3) Gnetoideae, genus Gnetum. Our knowledge of the Gnetales leaves much to be desired, but such facts as we possess would seem to indicate that this group is of special importance as foreshadowing, more than any other Gymnosperms, the Angiospermous type. In the more heterogeneous structure of the wood and in the possession of true vessels the Gnetales agree closely with the higher flowering plants. It is of interest to note that the leaves of Gnetum, while typically Dicotyledonous in appearance, possess a Gymnospermous character in the continuous and plate-like medullary rays of their vascular bundles. The presence of a perianth is a feature suggestive of an approach to the floral structure of Angiosperms; the prolongation of the integument furnishes the flowers with a substitute for a stigma and style. The genus Ephedra, with its prothallus and archegonia, which are similar to those of other Gymnosperms, may be safely regarded as the most primitive of the Gnetales. In Welwitschia also the megaspore is filled with prothallus-tissue, but single egg-cells take the place of archegonia. In certain species of Gnetum described by Karsten the megaspore contains a peripheral layer of protoplasm, in which scattered nuclei represent the female reproductive cells; in Gnetum Gnemon a similar state of things exists in the upper half of the megaspore, while the lower half agrees with the megaspore of Welwitschia in being full of prothallus-tissue, which serves merely as a reservoir of food. Lotsy has described the occurrence of special cells at the apex of the prothallus of Gnetum Gnemon, which he regards as imperfect archegonia (fig. 17, C, a); he suggests they may represent vestigial structures pointing back to some ancestral form beyond the limits of the present group. The Gnetales probably had a separate origin from the other Gymnosperms; they carry us nearer to the Angiosperms, but we have as yet no satisfactory evidence that they represent a stage in the direct line of Angiospermic evolution. It is not improbable that the three genera of this ancient phylum survive as types of a blindly-ending branch of the Gymnosperms; but be that as it may, it is in the Gnetales more than in any other Gymnosperms that we find features which help us to obtain a dim prospect of the lines along which the Angiosperms may have been evolved.
Ephedra.—This genus is the only member of the Gnetales represented in Europe. Its species, which are characteristic of warm temperate latitudes, are usually much-branched shrubs. The finer branches are green, and bear a close resemblance to the stems of Equisetum and to the slender twigs of Casuarina; the surface of the long internodes is marked by fine longitudinal ribs, and at the nodes are borne pairs of inconspicuous scale-leaves. The flowers are small, and borne on axillary shoots. A single male flower consists of an axis enclosed at the base by an inconspicuous perianth formed of two concrescent leaves and terminating in two, or as many as eight, shortly stalked or sessile anthers. The female flower is enveloped in a closely fitting sac-like investment, which must be regarded as a perianth; within this is an orthotropous ovule surrounded by a single integument prolonged upwards as a beak-like micropyle. The flower may be described as a bud bearing a pair of leaves which become fused and constitute a perianth, the apex of the shoot forming an ovule. In function the perianth may be compared with a unilocular ovary containing a single ovule; the projecting integument, which at the time of pollination secretes a drop of liquid, serves the same purpose as the style and stigma of an angiosperm. The megaspore
