Coniferales
CONIFERALES. Trees and shrubs characterized by a copious branching of the stem and frequently by a regular pyramidal form. Leaves simple, small, linear or short and scale-Tike, usually persisting for more than one year. Flowers monoecious or dioecious, unisexual, without a perianth, often in the form of cones, but never terminal on the main stem.
The plants usually included in the Coniferae constitute a less homogeneous class than the Cycadaceae. Some authors use the term Coniferae in a restricted sense as including those External genera which have the female flowers in the form of cones, features. the other genera, characterized by flowers of a different type, being placed in the Taxaceae, and often spoken of as Taxads.
In order to avoid confusion in the use of the term Coniferae, we may adopt as a class-designation the name Coniferales, including both the Coniferae using the term in a restricted sense and the Taxaceae. The most striking characteristic of the majority of the Coniferales is the regular manner of the monopodial branching and the pyramidal shape. Araucaria imbricata, the Monkey-puzzle tree, A. excelsa, the Norfolk Island pine, many pines and firs, cedars and other genera illustrate the pyramidal form. The mammoth redwood tree of California, Sequo ia (Wellingtonia) gigantea, which represents the tallest Gymnosperm, is a good example of the regular tapering main stem and narrow pyramidal form. The cypresses afford instances of tall and narrow trees similar in habit to Lombardy poplars. The common cypress (Cupressus sempervirens), as found wild in the mountains of Crete and Cyprus, is characterized by long and spreading branches, which give it a cedar-like habit. A pendulous or weeping habit is assumed by some conifers, e.g. Picea excelsa var. virgata represents a form in which the main branches attain a considerable horizontal extension, and trail themselves like snakes along the ground. Certain species of Pinus, the yews (Taxus) and some other genera grow as bushes, which in place of a main mast-like stem possess several repeatedly-branched leading shoots. The unfavourable conditions in Arctic regions have produced a dwarf form, in which the main shoots grow close to the ground. Artificially induced dwarfed plants of Pinus, Cupressus, Sciadopitys (umbrella pine) and other genera are commonly cultivated by the Japanese. The dying off of older branches and the vigorous growth of shoots nearer the apex of the stem produce a form of tree illustrated by the stone pine of the Mediterranean region (Pinus Pinea), which Turner has rendered familiar in his " Childe Harold's Pilgrimage " and other pictures of Italian scenery. Conifers are not infrequently seen in which a lateral branch has bent sharply upwards to take the place of the injured main trunk. An upward tendency of all the main lateral branches, known as fastigiation, is common in some species, producing wellmarked varieties, e.g. Cephalotaxus pedunculate var. fastigiata; this fastigiate habit may arise as a sport on a tree with spreading branches. Another departure from the normal is that in which the juvenile or seedling form of shoot persists in the adult tree; the numerous coniferous plants known as species of Retinospora are examples of this. The name Retinospora, therefore, does not stand for a true genus, but denotes persistent young forms of Juniperus, Thuja, Cupressus, etc., in which the small scaly leaves of ordinary species are replaced by the slender, needle-like leaves, which stand out more or less at right angles from the branches. The flat branchlets of Cupressus, Thuja (arbor vitae), Thujopsis dolabrata (Japanese arbor vitae) are characteristic of certain types of conifers; in some cases the horizontal extension of the branches induces a dorsiventral structure. A characteristic feature of the genus Agathis (Dammara) the Kauri pine of New Zealand, is the deciduous habit of the branches; these become detached from the main trunk leaving a well-defined absciss-surface, which appears as a depressed circular scar on the stem. A new genus of conifers, Tavwania, has recently been described from the island of Formosa; it is said to agree in habit with the Japanese Cryptomeria, but the cones appear to have a structure which distinguishes them from those of any other genus.
With a few exceptions conifers are evergreen, and retain the leaves for several years (10 years in Araucaria imbricata, 8 to 10 in Picea i .-, excelsa, 5 in Taxus baccata; in Pinus the needles usually fall in October of their third year). The larch (Larix) sheds its leaves in the autumn, in the Chinese larch ( Pseudolarix Kaempferi) the leaves turn a bright yellow colour before falling. In the swamp cypress (Taxodium distichum) the tree assumes a rich brown colour in the autumn, and sheds its leaves together with the branchlets which bear them; deciduous branches occur also in some other species, e.g. Sequoia sempervirens (redwood), Thuja occidentalis, etc. The leaves of conifers are characterized by their small size, e.g. the needle-form represented by Pinus, Cedrus, Larix, etc., the linear flat or angular leaves, appressed to the branches, of Thuja, Cupressus, Libocedrus, etc. The flat and comparatively broad leaves of Araucaria imbricata, A. Bidwillii, and some species of the southern genus Podocarpus are traversed by several parallel veins, as are also the still larger leaves of Agathis, which may reach a length of several inches. In addition to the foliage-leaves several genera also possess scale-leaves of various kinds, represented by budscales in Pinus, Picea, etc., which frequently persist for a time at the base of a young shoot which has pushed its way through the yielding cap of protecting scales, while in some conifers the bud-scales adhere together, and after being torn near the base are carried up by the growing axis as a thin brown cap. The cypresses, araucarias and some other genera have no true bud-scales; in some species, e.g. Araucaria Bidwillii, the occurrence of small foliage-leaves, which have functioned as bud-scales, at intervals on the shoots affords a measure of seasonal growth. The occurrence of long and short shoots is a characteristic feature of many conifers. In Pinus the needles occur in pairs, or in clusters of 3 or 5 at the apex of a small and inconspicuous short shoot of limited growth (spur), which is enclosed at its base by a few scale-leaves, and borne on a branch of unlimited growth in the axil of a scale-leaf. In the Californian Pinus monophylla each spur bears usually one needle, but two are not uncommon; it would seem that rudiments of two needles are always produced, but, as a rule, only one develops into a needle. In Sciadopitys similar spurs occur, each bearing a single needle, which in its grooved surface and in the possession of a double vascular bundle bears traces of an origin from two needle-leaves. A peculiarity of these leaves is the inverse orientation of the vascular tissue; each of the two veins has its phloem next the upper and the xylem towards the lower surface of the leaf; this unusual position of the xylem and phloem may be explained by regarding the needle of Sciadopitys as being composed of a pair of leaves borne on a short axillary shoot and fused by their margins (fig. 15, A). Long and short shoots occur also in Cedrus and Larix, but in these genera the spurs are longer and stouter, and are not shed with the leaves; this kind of short shoot, by accelerated apical growth, often passes into the conditio* of a long shoot on which the leaves are scattered and separated by comparatively long internodes, instead of being crowded into tufts such as are borne on the ends of the spurs. In the geniis PhyUocladus (New Zealand, etc.) there are no green foliage-leaves, but in their place flattened branches (phylloclades) borne in the axils of small scaleleaves. The cotyledons are often two in number, but sometimes (e.g. Pinus) as many as fifteen; these leaves are usually succeeded by foliage-leaves in the form of delicate spreading needles, and these primordial leaves are followed, sooner or later, by the adult type of leaf, except in Retinosporas, which retain the juvenile foliage. In addition to the first foliage-leaves and the adult type of leaf, there are often produced leaves which are intermediate both in shape and structure between the seedling and adult foliage. Dimorphism or heterophylly is fairly common. One of the best known examples is the Chinese juniper (Juniperus chinensis), in which branches with spinous leaves, longer and more spreading than the ordinary adult leaf, are often found associated with the normal type of branch. In some cases, e.g. Sequoia sempervirens, the fertile branches bear leaves which are less spreading than those on the vegetative shoots. Certain species of the southern hemisphere genus Dacrydium afford particularly striking instances of heterophylly, e.g. D. Kirkii of New Zealand, in which some branches bear small and appressed leaves, while in others the leaves are much longer and more spreading. A wellknown fossil conifer from Triassic strata Voltzia heterophylla also illustrates a marked dissimilarity in the leaves of the same shoot. The variation in leaf-form and the tendency of leaves to arrange themselves in various ways on different branches of the same plant are features which it is important to bear in mind in the identification of fossil conifers. In this connexion we may note the striking resemblance between some of the New Zealand Alpine Veronicas, e.g. Veronica Hectori, V.cupressoides,etc. (alsoPolycladuscupressinus, a Composite), and some of the cypresses and other conifers with small appressed leaves. The long linear leaves of some species of Podocarpus, in which the lamina is traversed by a single vein, recall the pinnae of Cycas; the branches of some Dacrydiums and other forms closely resemble those of lycopods; these superficial resemblances, both between different genera of conifers and between conifers and other plants, coupled with the usual occurrence of fossil coniferous twigs without cones attached to them, render the determination of extinct types a very unsatisfactory and frequently an impossible task.
A typical male flower consists of a central axis bearing numerous spirally-arranged sporophylls (stamens), each of which consists of a slender stalk (filament) terminating distally in a more Fi ower s. or less prominent knob or triangular scale, and bearing two or more pollen-sacs (microsporangia) on its lower surface. The pollen-grains of some genera (e.g. Pinus) are furnished with bladderlike extensions of the outer wall, which serve as aids to wind -dispersal. The stamens of Araucaria and Agathis are peculiar in bearing several long and narrow free pollen-sacs; these may be compared with the sporangiophores of the horsetails (Equisetum) ; in Taxus (yew) the filament is attached to the centre of a large circular distal expansion, which bears several pollen-sacs on its under surface. In the conifers proper the female reproductive organs have the form of cones, which may be styled flowers or inflorescences according to different interpretations of their morphology. In the Taxaceae the flowers have a simpler structure. The female flowers of the Abietineae may be taken as representing a common type. A pine cone reaches maturity in two years; a single year suffices for the full development in Larix and several other genera. The axis of the cone bears numerous spirally disposed flat scales (cone-scales), each of which, if examined in a young cone, is found to be double, and to consist of a lower and an upper portion. The latter is a thin flat scale bearing a median ridge or keel (e.g. Abies), on each side of which is situated an inverted ovule, consisting of a nucellus surrounded by a single integument. As the cone grows in size and becomes woody the lower half of the cone-scale, which we may call the carpellary scale, may remain small, and is so far outgrown by the upper half (seminiferous scale) that it is hardly recognizable in the mature cone. In many species of Abies (e.g. Abies pectinata, etc.) the ripe cone differs from those of Pinus, Picea and Cedrus in the large size of the carpellary scales, which project as conspicuous thin appendages beyond the distal margins of the broader and more woody seminiferous scales; the long carpellary scale is a prominent feature also in the cone of the Douglas pine (Pseudotsuga Douglasii). The female flowers (cones) vary considerably in size; the largest are the more or less spherical cones of Araucaria a single cone of A. imbricata may produce as many as 300 seeds, one seed to each fertile cone-scale and the long pendent female /lower.
cones, i to 2 ft. in length, of the sugar pine of California (Pinus Lambertw.no) and other species. Smaller cones, less than an inch long, occur in the larch, Athrotaxis (Tasmania), Fitzroya (Patagonia and Tasmania), etc. In the Taxodieae and Araucarieae the cones are similar in appearance to those of the Abietineae, but they differ in the fact that the scales appear to be single, even in the young condition ; each cone-scale in a genus of the Taxodiinae (Sequoia, etc.) bears several seeds, while in the Araucariinae (Araucaria and Agathis) each scale has one seed. The Cupressineae have cones composed of a few scales arranged in alternate whorls; each scale bears two or more seeds, and shows no external sign of being composed of two distinct pbrtions. In the junipers the scales become fleshy as the seeds ripen, and the individual scales fuse together in the form of a berry. The female flowers of the Taxaceae assume another form ; in Microcachrys (Tasmania) the reproductive structures are spirally disposed, and form small globular cones made up of red fleshy scales, to each of which is attached a single ovule enclosed by an integument and partially invested by an a rill us; in Dacrydium the carpellary leaves are very similar to the foliage leaves each bears one ovule with two integuments, the outer of which constitutes an arillus. Finally in the yew, as a type of the family Taxeae, the ovules occur singly at the apex of a lateral branch, enclosed when ripe by a conspicuous red or yellow fleshy arillus, which serves as an attraction to animals, and thus aids in the dispersal of the seeds.
It is important to draw attention to some structural features exhibited by certain cone-scales, in which there is no external sign indicative of the presence of a carpellary and a seminiferous Morpho- scale, j n Araucaria Cookii and some allied species each scale has a small pointed projection from its upper face near the distal end ; the scales of Cunninghamia (China) are characterized by a somewhat ragged membranous projection extending across the upper face between the seeds and the distal end of the scale; in the scales of Athrotaxis (Tasmania) a prominent rounded ridge occupies a corresponding position. These projections and ridges may be homologous with the seminiferous scale of the pines, firs, cedars, etc. The simplest interpretation of the cone of the Abietineae is that which regards it as a flower consisting of an axis bearing several open carpels, which in the adult cone may be very small or large and prominent, the scale bearing the ovules being regarded as a placental outgrowth from the flat and open carpel. In Araucaria the cone-scale is regarded as consisting of a flat carpel, of which the placenta has not grown out into the scale-like structure. The seminiferous scale of Pinus, etc., is also spoken of sometimes as a ligular outgrowth from the carpellary leaf. Robert Brown was the first to gjve a clear description of the morphology of the Abietineous cone in which carpels bear naked ovules ; he recognized gymnospermy as an important distinguishing feature in conifers as well as in cycads. Another view is to regard the cone as an inflorescence, each carpellary scale being a bract bearing in its axil a shoot the axis of which has not been developed; the seminiferous scale is believed to represent either a single leaf or a fused pair of leaves belonging to the partially suppressed axillary shoot. In 1869 van Tieghem laid stress on anatomical evidence as a key to the morphology of the cone-scales; he drew attention to the fact that the collateral vascular bundles of the seminiferous scale are inversely orientated as compared with those of the carpellary scale ; in the latter the xylem of each bundle is next the upper surface, while in the seminiferous scale the phloem occupies that position. The conclusion drawn from this was that the seminiferous scale (fig. 15,8, Sc) is the first and only leaf of an axillary shoot (b) borne on that side of the shoot, the axis of which is suppressed, opposite the subtending bract (fig. 15, A, B, C, Br). Another view is to apply to the seminiferous scale an explanation similar to that suggested by yon Mohl in the case of the double needle of Sciadopitys, and to consider the seed-bearing scale as being made up of a pair of leaves (fig. 15, A, a, a) of an axillary shoot (b) fused into one by their posterior margins (fig. 15, A). The latter view receives support from abnormal cones in which carpellary scales subtend axillary shoots, of which the first two leaves (fig. 15, C, / l , I 1 ) are often harder and browner than the others; ' forms have been described transitional between axillary shoots, in which the leaves are separate, and others in which two of the leaves are more or less completely fused. In a young cone the seminiferous scale appears as a hump of tissue at the base or in the axil of the carpellary scale, but Celakovsky, a strong supporter of the axillary-bud theory, attaches little or no importance to this kind of evidence, regarding the present manner of development as being merely an example of a short cut adopted in the course of evolution, and replacing the original production of a branch in the axil of each carpellary scale. Eichler, one of the chief supporters of the simpler view, does not recognize in the inverse orientation of the vascular bundles an argument in support of the axillary-bud theory, but points out that the seminiferous scale, being an outgrowth from the surface of the carpellary scale, would, like outgrowths from an ordinary leaf, naturally have its bundles inversely orientated. In such cone-scales as show little or no external indication of being double in origin, e.g. Araucaria (fig. 15, D) Sequoia, etc., there are always two sets of bundles; the upper set, having the phloem uppermost, as in the seminiferous scale of Abies or Pinus, are regarded as belonging to the outgrowth from the carpellary scale and specially developed to supply the ovules. Monstrous cones are fairly common; these in some instances lend support to the axillary-bud theory, and it has been said that this theory owes its existence to evidence furnished by abnormal cones. It is_ difficult to estimate the value of abnormalities as evidence bearing on morphological interpretation; the chief danger lies perhaps in attaching undue weight to them, but there is also a risk of minimizing their importance. Monstrosities at least demonstrate possible lines of development, but when the abnormal forms of growth in various directions are fairly evenly balanced, trustworthy deductions become difficult. The occurrence of buds in the axils of carpellary scales may, however, simply mean that buds, which are (C and D after Worsdell.)
FIG. 15. Diagrammatic treatment of:
A, Double needle of Sciadopitys (a, a, leaves; b, shoot ; Br, bract).
B, seminiferous scale as leaf of axillary shoot (b, shoot ; Sc, semi- niferous scale ; Br, bract).
C, seminiferous scale as fused pair of leaves (P, P, I 3 , first, second and third leaves; b, shoot; Br, bract), D, cone-scale of Araucaria (n, nucellus; i, integument; *, xylem).
usually undeveloped in the axils of sporophylls, occasionally afford evidence of their existence. Some monstrous cones lend no support to the axillary-bud theory. In Larix the axis of the cone often continues its growth ; similarly in Cephalotaxus the cones are often proliferous. (In rare cases the proliferated portion produces male flowers in the leaf-axils.) In Larix the carpellary scale may become leafy, and the seminiferous scale may disappear. Androgynous cones may be produced, as in the cone of Pinus rigida (fig. 16), in which the lower part bears stamens and the upper portion carpellary and seminiferous scales. An interesting case has been figured by Masters, in which scales of a cone of Cupressus Lawsoniana bear ovules on the upper surface and stamens on the lower face. One argument that has been adduced in support of the axillary bud theory is derived from the Palaeozoic type Cordaites, in which each ovule occurs on an axis borne in the axil of a bract. The whole question is still unsolved, and perhaps insoluble. It may be that the interpretation of the female cone of the Abietineae as an inflorescence, which finds favour with many botanists, cannot be applied to the cones of Agathis and Araucaria. Without expressing any decided opinion as to the morphology of the double cone-scale of the Abietineae, preference may be felt in favour of regarding the cone-scale of the Araucarieae as a simple carpellary leaf bearing a single ovule. A discussion of this question may be found in a paper on the Araucarieae by Seward and Ford, published in the Transactions of the Royal Society of London (1906). Cordaites is an extinct type normal Lone ot which in certain respects resembles Ginkgo, cycads ft?**** and the Araucarieae, but its agreement with true (After Masters.) conifers is probably too remote to justify our attributing much weight to the bearing of the morphology of its female flowers on the interpretation of that of the Coniferae. The greater simplicity of the Eichler theory may prejudice us in its favour; but, on the other hand, the arguments advanced in favour of the axillary-bud theories are perhaps not sufficiently cogent to lead us to accept an explanation based chiefly on the uncertain evidence of monstrosities.
A pollen-grain when first formed from its mother-cell consists of a single cell; in this condition it may be carried to the nucellus of the ovule (e.g. Taxus, Cupressus, etc.), or more usually . (Pinus, Larix, etc.) it reaches maturity before the dehis- mlcr0 ' cence of the microsporangium. The nucleus of the a a a mella . microspore divides and gives rise to a small cell within spores the large cell, a second small cell is then produced; this is the structure of the ripe pollen-grain in some conifers (Taxus, etc.). The large cell grows out as a pollen-tube; the second of the two small cells (body-cell) wanders into the tube, followed by the nucleus of the first small cell (stalk-cell). In Taxus the body-cell eventually divides into two, in which the products of division are of unequal size, the larger constituting the male generative cell, which fuses with the nucleus of the egg-cell. In Juniperus the products of division of the FIG. 16 Ab- body-cell are equal, and both function as male generative cells. In the Abietineae cell-formation in the pollen-grain is carried farther. Three small cells occur inside the cavity of the microspore; two of them collapse and the third divides into two, forming a stalk-cell and a larger body-cell. The latter ultimately divides in the apex of the pollen-tube into two non-motile generative cells. Evidence has lately been adduced of the existence of numerous nuclei in the pollen-tubes of the Araucarieae, and it seems probable that in this as in several other respects this family is distinguished from other members of the Coniferales. The precise method of fertilization in the Scots Pine was followed by V. H. Blackman, who also succeeded in showing that the nuclei of the sporophyte generation contain twice as many chromosomes as the nuclei of the gametophyte. Other observers have in recent years demonstrated a similar relation in other genera between the number of chromosomes in the nuclei of the two generations. The ovule is usually surrounded by one integument, which projects beyond the tip of the nucellus as a wide-open Ipbed funnel, which at the time of pollination folds inwards, and so assists in bringing the pollen-grains on to the nucellus. In some _ conifers (e.g. Taxus, Cephalotaxus, Dacrydium, etc.) the ordinary integument is partially enclosed by an arillus or second integument. It is held by some botanists (Celakovsky) that the seminiferous scale of the Abietineae is homologous with the arillus or second integument of the Taxaceae, but this view is too strained to gain general acceptance. In Araucaria and Saxegothaea the nucellus itself projects beyond the open micropyle and receives the pollen-grains direct. During the growth of the cell which forms the megaspore the greater part of the nucellus is absorbed, except the apical portion, which persists as a cone above the megaspore ; the partial disorganization of some of the cells in the centre of the nucellar cone forms an irregular cavity, which may be compared with the larger pollen-chamber of Ginkgo and the cycads. In each ovule one megaspore comes to maturity, but, exceptionally, two may be present (e.g. Pinus sylvestris). It has been shown by Lawson that in Sequoia sempervirens (Annals of Botany, 1904) and by other workers in the genera that several megaspores may attain a fairly large size in one prothallus. The megaspore becomes filled with tissue (prothallus), and from some of the superficial cells archegonia are produced, usually three to five in number, but in rare cases ten to twenty or even sixty may be present. In the genus Sequoia there may be as many as sixty archegonia (Arnold! and Lawson) in one megaspore ; these occur either separately or in some parts of the prothallus they may form groups as in the Cupressineae ; they are scattered through the prothallus instead of being confined to the apical region as in the majority of conifers. Similarly in the Araucarieae and in Widdringtonia the archegonia are numerous and scattered and often sunk in the prothallus tissue. In Libocedrus decurrens (Cupressineae) Lawson describes the archegonia as varying in number from 6 to 2\(Annals oj 'Botany xxi.,1907). An archegonium consists of a large oval egg-cell surmounted by a short neck composed of one or more tiers of cells, six to eight cells in each tier. Before fertilization the nucleus of the egg-cell divides and cuts off a ventral canal-cell; this cell may represent a second egg-cell. The egg-cells of the archegonia may be in lateral contact (e.g. Cupressineae) or separated from one another by a few cells of the prothallus, each ovum being immediately surrounded by a layer of cells distinguished by their granular contents and large nuclei. During the development of the egg-cell, food material is transferred from these cells through the pitted wall of the ovum. The tissue at the apex of the megaspore grows slightly above the level of the archegonia, so that the latter come to Re in a shallow depression. In the process of fertilization the two male generative nuclei, accompanied by the pollen-tube nucleus and that of the stalk-cell, pass through an open pit at the apex of the pollen-tube into the protoplasm of the ovum. After fertilization the nucleus of the egg divides, the first stages of karyokinesis being apparent even before complete fusion of the male and female nuclei has occurred. The result of this is the production of four nuclei, which eventually take up a position at the bottom of the ovum and become separated from one another by vertical cellwalls; these nuclei divide again, and finally three tiers of cells are produced, four in each tier. In the Abietineae the cells of the middle tier elongate and push the lowest tier deeper into the endosperm; the cells of the bottom tier may remain in lateral contact and produce together one embryo, or they may separate (Pinus, Juniperus, etc.) and form four potential embryos. The ripe albuminous seed contains a single embryo with two or more cotyledons. The seeds of many conifers are provided with large thin wings, consisting in some genera (e.g. Pinus) of the upper cell-layers of the seminiferous scale, which have become detached and, in some cases, adhere loosely to the seed as a thin membrane ; the loose attachment may be of use to the seeds when they are blown against the branches of trees, in enabling them to fall away from the wing and drop to the ground. The seeds of some genera depend on animals for dispersal, the carpellary scale ( Microcachrys) or the outer integument being brightly coloured and attractive. In some Abietineae (e.g. Pinus and Picea) in which the cone-scales persist for some time after the seeds are ripe the cones hang down and so facilitate the fall of the seeds; in Cedrus, Araucaria and Abies the scales become detached and fall with the seeds, leaving the bare vertical axis of the cone on the tree. In all cases, except some species of Araucaria (sect. Colymbea) the germination is epigean. The seedling plants of some Conifers (e.g. Araucaria imbricata) are characterized by a carrot-shaped hypocotyl, which doubtless serves as a food-reservoir.
The roots of many conifers possess a narrow band of primary xylem-tracheids with a group of narrow spiral protoxylem-elements at each end (diarch). A striking feature in the roots of several genera, excluding the Abietineae, is the occur- Anatomy. rence of thick and somewhat irregular bands of thickening on the cell-walls of the cortical layer next to the endodermis. These bands, which may serve to strengthen the central cylinder, have been compared with the netting surrounding the delicate wall of an inflated balloon. It is not always easy to distinguish a root from a stem; in some cases (e.g. Sequoia) the primary tetrarch structure is easily identified in the centre of an old root, but in other cases the primary elements are very difficult to recognize. The sudden termination of the secondary tracheids against the pith-cells may afford evidence of root-structure as distinct from stem-structure, in which the radial rows of secondary tracheids pass into the irregularly-arraneed primary elements next the pith. The annual rings in a root are often less clearly marked than in the stem, and the xylem-elements are frequently larger and thinner. The primary vascular bundles in a young conifer stem are collateral, and, like those of a Dicotyledon, they are arranged in a circle round a central pith and enclosed by a common endodermis. It is in the nature of the secondary xyfem that the Coniferales are most readily distinguished from the Dicotyledons and Cycadaceae; the wood is homogeneous in structure, consisting almost entirely of tracheids with circular or polygonal bordered pits on the radial walls, more particularly in the late summer wood. In many genera xylem-parenchyma is present, but never in great abundance. A few Dicotyledons, e.g. Drimys (Magnoliaceae) closely resemble conifers in the homogeneous character of the wood, but in most cases the presence of large spring vessels, wood-fibres and abundant parenchyma affords an obvious distinguishing feature.
The abundance of petrified coniferous wood in rocks of various ages has led many botanists to investigate the structure of modern genera with a view to determining how far anatomical characters may be used as evidence of generic distinctions. There are a few well-marked types of wood which serve as convenient standards of comparison, but these cannot be used except in a few cases to distinguish individual genera. The genus Pinus serves as an illustration of wood of a distinct type characterized by the absence of xylemparenchyma, except such as is associated with the numerous resincanals that occur abundantly in the wood, cortex and medullary rays; the medullary rays are composed of parenchyma and of horizontal tracheids with irregular ingrowths from their walls. In a radial section of a pine stem each ray is seen to consist in the median part of a few rows of parenchymatous cells with large oval simple pits in their walls, accompanied above and below by horizontal tracheids with bordered pits. The pits in the radial walls of the ordinary xylem-tracheids occur in a single row or in a double row, of which the pits are not in contact, and those of the two rows are placed on the same level. The medullary rays usually consist of a single tier of cells, but in the Pinus type of wood broader medullary rays also occur and are traversed by horizontal resin-canals. In the wood of Cypressus, Cedrus, Abies and several other genera, parenchymatous cells occur in association with the xylem-tracheids and take the place of the resin-canals of other types. In the Araucarian type of wood (Araucaria and Agathis) the bordered pits, which occur in two or three rows on the radial walls of the tracheids, are in mutual contact and polygonal in shape, the pits of the different rows are alternate and not on the same level ; in this type of wood the annual rings are often much less distinct than in Cupressus, Pinus and other genera. In Taxus, Torreya (California and theFar East) and Cephalotaxus the absence of resin-canals and the presence of spiral thickeningbands on the tracheids constitute well-marked characteristics. An examination of the wood of branches, stems and roots of the same species or individual usually reveals a fairly wide variation in some of the characters, such as the abundance and size of the medullary rays, the size and arrangement of pits, the presence of wood-parenchyma characters to which undue importance has often been attached in systematic anatomical work. The phloem consists of sieve-tubes, with pitted areas on the lateral as well as on the inclined terminal walls, phloem-parenchyma and, in some genera, fibres. In the Abietineae the phloem consists of parenchyma and sieve-tubes only, but in most other forms tangential rows of fibres occur in regular alternation with the parenchyma and sieve-tubes. The characteristic companion-cells of Angiosperms are represented by phloem-parenchyma cells with albuminous contents; other parenchymatous elements of the bast contain starch or crystals of calcium oxalate. When tracheids occur in the medullary rays of the xylem these are replaced in the phloem-region by irregular parenchymatous cells known as albuminous cells. Resin-canals, which occur abundantly in the xylem, phloem or cortex, are not found in the wood of the yew. Cephalotaxus (Taxeae) is also peculiar in having resincanals in the pith (cf. Ginkgo). One form of Cephalotaxus is characterized by the presence of short tracheids in the pith, in shape like ordinary parenchyma, but in the possession of bordered pits and lignified walls agreeing with ordinary xylem-tracheids; it is probable that these short tracheids serve as reservoirs for storing rather than for conducting water. The vascular bundle entering the stem from a leaf with a single vein passes by a more or less direct course into the 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, etc.) the leaf-trace leaves the stele of the stem as a single bundle which spJits 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, etc. 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 coniferpus leaves; these elements assume two distinct forms (i) 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, etc. 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, etc., 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, etc., a single canal occurs below the midrib; in Larix, Abies, etc., 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 Dlstrlbu- ant ' P 6 " 611 " 21 ' 6 beyond the northern limit of dicotyledonous trees. Wide areas are often exclusively occupied by 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, etc. In north Siberia Pinus Cembra (Cembra or Arolla Pine) has a wide range; also Abies 6i>;a(Siberian silver fir), Larix sibirica and Juniperus Sabina(sa.v'm). 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, etc. 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), Taxpdium distichum (swamp cypress), Pinus rigida (pitch pine), P. mitis (yellow pine), P. taeda,P. palustris, etc. On the west side of the American continent conifers play a still more striking r61e ; 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, etc. 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,Cryptpmeria japonica, Cephalotaxus, species of A bies, Larix, Thujopsis, Sciadopitys verticillata, Pseudolarix Kaempferi, etc. In the Himalaya occur Cedrus deodara, Taxus, species of Cupressus, Pinus excelsa, Abies Webbiana, etc. 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. milaniiana 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. Atkrotaxis 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.
Note - this article incorporates content from Encyclopaedia Britannica, Eleventh Edition, (1910-1911)
