The Genus Pinus

THE CONE. Plate IV.

The cone of Pinus shows great differences of color, form and tissue; these are useful for specific and sectional distinctions, while the gradual change from the primitive conditions of the Cembrae to the elaborate form, structure and mode of dissemination of some serotinous species are obvious evidence of an evolution among the species of remarkable taxonomic range. A form new among Coniferae appears, the oblique cone, and a new condition, the serotinous cone, both appearing at first alone and, finally, in constant association.

COLOR OF THE CONE.

With few exceptions the color of the ripe cone may be classified under one of the following shades of brown or yellow.

These colors may be paler or deeper. They may be obscured by a fuscous shade or may be modified by a dull or lustrous surface. The presence of two or more of these shades in a single species and the inherent difficulties of color description lessen the value of the character. Nevertheless certain allied species, such as P. nigra and Thunbergii, or P. densiflora and Massoniana, may be distinguished by the prevalent difference in the color of their cones.

DIMENSIONS OF THE CONE.

The cone is small, medium or large in different species, but varies greatly under the influences of environment or of individual peculiarities. The character possesses relative value only, for great variation is possible in the same locality and even on the same tree.

THE PEDUNCLE.

All conelets are pedunculate, but in some species the peduncle, even when long (patula), may become overgrown and concealed by the basal scales of the ripe cone. Articulation usually takes place between the peduncle and the branch, sometimes with the loss of a few basal scales which remain temporarily on the tree (ponderosa, palustris, etc.). With P. Nelsonii, and to a less degree with P. Armandi, there is articulation between the cone and its peduncle.

There are several species bearing persistent cones with no articulation. This condition appears in other genera, such as Larix and Picea, but without obvious significance. In Pinus, however, the gradual appearance of the persistent cone, for it is rare, common, prevalent or invariable in different species, and its essential association with the serotinous cone, suggest an evolution toward a definite end.

THE UMBO.

The exposed part of the scale of the conelet is the umbo of the ripe cone, a small definite area representing the earlier part of the biennial growth of the cone. The position of the umbo on the apophysis is the basis of Koehne's subdivision of the section Haploxylon.

Two other characters assist in establishing these subsections—the conelet, unarmed in Cembra, armed in Paracembra—the pits of the ray-cells of the wood, large in Cembra, small in Paracembra.

THE APOPHYSIS.

The apophysis represents the later and larger growth of the cone-scale. With a terminal umbo the margin of the apophysis is free and may be rounded (fig. 49) or may taper to a blunt point (fig. 52), and any extension of the scale is a terminal extension. With the dorsal umbo all sides of the apophysis are confined between other apophyses, and any extension is a dorsal thickening of the apophysis or a dorsal protuberance. The outline of an apophysis with a dorsal umbo is quadrangular, or it is irregularly pentagonal or hexagonal, the different forms depending on the arrangement of the contiguous scales, whether of definite or indefinite phyllotactic order, a distinction to be considered later.

The two positions of the umbo result from the relative growth of the dorsal and ventral surfaces of the cone-scale. With the terminal umbo the growth of both surfaces is uniform, with the dorsal umbo the growth is unequal. A true terminal umbo rests on the surface of the underlying scale, although several species with terminal umbos show the first stages of the dorsal umbo. The umbo of P. Lambertiana or of P. flexilis does not touch the surface of the scale below, and a small portion of the under side of the apophysis is brought into view on the closed cone. The cone of P. albicaulis (Plate VIII, fig. 90) shows all degrees of development between a terminal umbo near the apex of the cone and a dorsal umbo near its base.

The growth of the apophysis may be limited and constant (strobus, echinata, etc.) or exceedingly variable, ranging from a slight thickness to a long protuberance (pseudostrobus, montana, etc.). The protuberance is usually reflexed from the unequal growth of the two surfaces. With the terminal umbo the protuberance lengthens the scale, with the dorsal umbo it thickens the scale. It is sometimes a specific character (ayacahuite, longifolia) appearing on all cones of the species, sometimes a varietal form, associated in the same species with an unprolonged apophysis (sylvestris, montana).

On different parts of the same cone, base, centre or apex, the dimensions of the apophyses differ, but at each level the scales may be uniform on all sides of the cone. That is to say, the cone is symmetrical with reference to any plane passing through its axis. This, the symmetrical cone, is characteristic of all other genera of the Abietineae, and is invariable among the Soft Pines and in many Hard Pines (figs. 47, 48, 52, 54). But among the Hard Pines there is gradually developed a new form of cone with smaller flatter apophyses on the anterior, and larger thicker apophyses on the posterior surface. This is the peculiar oblique cone of Pinus (figs. 50, 51, 53), symmetrical with reference to one plane only, which includes the axis of both cone and branch. The oblique cone is a gradual development among the Hard Pines; in some species it is associated as a varietal form with the symmetrical cone, and finally, in some serotinous species, it is the constant form.

THE OBLIQUE CONE.

When the oblique cone is merely a varietal form (halepensis, etc.), it gives the impression of an accident, resulting from the reflexed position of the cone and the consequent greater development of the scales receiving a greater amount of light and air. But with the serotinous cones (radiata, attenuata), the advantages of this form become apparent. The cones of these species are in crowded nodal clusters, reflexed against the branch (fig. 50). The inner, anterior scales are perfectly protected by their position, while the outer, posterior scales are exposed to the weather. These last only are very thick; that is to say, there is an economical distribution of protective tissue, with the greatest amount where it is most needed. The oblique form is peculiarly adapted for a cone destined to remain on the tree for twenty years or more and to preserve its seeds unimpaired. Like the persistent cone, the oblique cone finds in association with the serotinous cone a definite reason for existence.

PHYLLOTAXIS. Plate V.

There is an obvious difference between the cones of the two sections of the genus. Those of the Soft Pines (figs. 55, 56) have larger and fewer scales, those of the Hard Pines (figs. 57, 58) have more numerous and smaller scales, in proportion to the size of the cone. The former condition represents a lower, the latter condition represents a higher, order of phyllotaxis.

DEFINITE PHYLLOTAXIS.

On a cylindrical axis with scales of the same size, the spiral arrangement would appear as in fig. 62, where the scales are quadrangular and any four adjacent scales are in mutual contact at their sides or angles. These four scales lie on four obvious secondary spirals (fig. 59, a-a, b-b, c-c, d-d). According to the phyllotactic order of the scales these may be the spirals of 2, 3, 5, 8 or of 3, 5, 8, 13 or of 5, 8, 13, 21 etc., etc., from which combinations the primary spiral, on which the scales are inserted on the cone-axis, can be easily deduced. Four quadrangular scales in mutual contact represent the condition of definite phyllotaxis. If the cone is conical, definite phyllotaxis would be possible among all the scales only when the size of the scales diminishes in equal measure with the gradual diminution of the cone's diameter. Such a hypothetical cone is shown in fig. 61.

INDEFINITE PHYLLOTAXIS.

On an imaginary cone of conical form and with scales of equal size throughout, there must be more scales about the base than about the apex of the cone. The phyllotactic conditions must differ, and the obvious spirals, in passing from base to apex, must undergo readjustment. If the scales at the base are in definite phyllotactic order and those at the apex are in the next lower order, it is evident that intermediate scales, in the gradual change from one condition to the other, must represent different conditions of indefinite phyllotaxis, while those in a central position on the cone may belong equally to either of two orders.

A Pine cone is never absolutely cylindrical nor do its scales vary in size proportionately to the change of diameter. Most of the scales of a cone are in indefinite phyllotactic relation, while definite phyllotaxis is found only at points on the cone.

As an extreme illustration, the cone of P. pinaster (fig. 60) shows four mutually contiguous quadrangular apophyses at (a), lying on the obvious spirals 5, 8, 13, 21, at (b) four similar apophyses on the spirals 3, 5, 8, 13, and at (c) four others on the spirals 2, 3, 5, 8. Between these three points are apophyses of irregular pentagonal or hexagonal outline, with three scales only in mutual contact (figs. 63, 64). Such are the majority of the scales of the cone and represent more or less indefinite conditions of phyllotaxis.

The cones of Hard Pines, by reason of relatively more and smaller scales and of a more conical form, attain a higher phyllotaxis and a more complex condition, two or even three orders being represented on a single cone; while the cones of Soft Pines, by reason of relatively fewer and larger scales and a more cylindrical form, are of lower phyllotaxis, with one order only more or less definitely presented. Therefore phyllotaxis furnishes another distinction between the two sections of the genus, but its further employment is exceedingly restricted on account of the constant repetition of the same orders among the species.

THE CONE-TISSUES. Plate VI.

The axis of the cone is a woody shell, enclosing a wide pith and covered by a thick cortex traversed by resin-ducts. By removing the scales and cortex from the axis (fig. 65) the wood is seen to be in sinuous strands uniting above and below fusiform openings, the points of insertion of the cone-scales. From the wood, at each insertion, three stout strands enter the scale, dividing and subdividing into smaller tapering strands whose delicate tips converge toward the umbo. Fig. 70 represents a magnified cross-section of half the cone-scale of P. Greggii; at (a) is a compact dorsal plate of bast cells; at (e) is a ventral plate of the same tissue but of less amount; at (b) is the softer brown tissue enclosing the wood-strands (d, d) (the last much more magnified in fig. 69) and the resin-ducts (e, e).

WOOD STRANDS.

The wood-strands, forming the axis of the cone, differ in tenacity in the two sections of the genus. Those of the Soft Pines are easily pulled apart by the fingers, those of the Hard Pines are tougher in various degrees and cannot be torn apart without the aid of a tool. This difference is correlated with differences in other tissues, all of them combining in a gradual change from a cone of soft yielding texture to one of great hardness and durability.

If a cone scale of P. ayacahuite is stripped of its brown and bast tissues (fig. 66) and is immersed in water and subsequently dried, there is at first a flexion toward the cone-axis (fig. 67) and then away from it (fig. 68). The wood-strands are hygroscopic and coöperate with the bast tissues in opening and closing the cone. This appears to be true of all species excepting the three species of the Cembrae, whose strands are so small and weak that they are not obviously affected by hygrometric changes.

BAST TISSUE.

With the exception of the three species of the Cembrae the inner part of the cone-scales is protected by sclerenchymatous cells forming hard dorsal and ventral plates (fig. 70, a, c). In Soft Pines these cells are subordinate to the more numerous parenchymatous cells, but in Hard Pines the sclerenchyma increases in amount until, among the serotinous species, it is the predominating tissue of the cone-scale, giving to these cones their remarkable strength and durability.

This bast tissue is hygroscopic and, with its greater thickness on the dorsal surface, there is a much greater strain on that side of the scale, tending to force the scales apart when they are ripe and dry, and subsequently closing and opening the cone on rainy and sunny days.

The cone, during the second season's growth, is completely closed, its scales adhering together with more or less tenacity. In most species the hygroscopic energy of the scales is sufficient to open the cone under the dry condition of its maturity, but with several species the adhesion is so persistent that some of the cones remain closed for many years. These are the peculiar serotinous cones of the genus.

THE SEROTINOUS CONE.

As an illustration of the area to which the adhesion is confined, a section may be sawed from a cone of P. attenuata (fig. 71). The axis and the scales that have been severed from their apophyses (b) can be easily pushed out of the annulus (a), which is composed wholly of apophyses so firmly adherent that they will successfully resist a strong effort to break them apart. When immersed in boiling water, however, the ring falls to pieces. An examination of these pieces discovers adhesion only on a narrow ventral border under the apophysis and on a corresponding dorsal border back of the apophysis. The rest of the scale is not adherent, so that the seed is free to fall at the opening of the cone.

The serotinous cone is a gradual development, wanting in most species, rare in a few, less or more frequent in others. A similar evolution of the persistent cone, of the oblique cone and of the cone-tissues has been already discussed. All these progressive characters culminate in mutual association in P. radiata and its allies. The result is a highly specialized fruit that should convey taxonomic significance of some kind.

With all serotinous species that I have seen, some of the trees open their cones at maturity, others at indefinite intervals. That is to say, the seed of a prolific year is not at the mercy of a single, perhaps unfavorable season. The chances of successful germination are much increased by the intermittent seed-release peculiar to these Pines. Such a method of dissemination must accrue to the advantage of a species. In other words, this intermittent dissemination and the oblique form of cone with its perfected tissues all mark the highest development of the genus.

THE SEED. Plate VI. Figs. 72-79.

The seed of Pinus contains an embryo, with the cotyledons clearly defined, embedded in albumen, which is protected by a bony testa with an external membranous spermoderm, produced, in most species, into an effective wing. While the seed of other genera of the Abietineae shows no striking difference among the species, that of Pinus is remarkably variable, presenting alike the most primitive and the most elaborate forms among the Conifers. These differences are valuable for the segregation of kindred species and for some specific distinctions.

WINGLESS SEEDS.

With wingless seeds the main distinction is found in the spermoderm, which is entire in one species only, P. koraiensis. In P. cembra it is wanting on the ventral surface of the nut, but on the dorsal surface, it is adnate partly to the nut, partly to the cone-scale. The nut of P. albicaulis and that of P. cembroides are quite bare of membranous cover. The spermoderm of P. flexilis is reduced to a marginal border, slightly produced into a rudimentary wing adnate to the nut.

THE ADNATE WING.

In P. strobus, longifolia and their allies and in P. Balfouriana the spermoderm is prolonged into an effective wing-blade from a marginal adnate base like that of P. flexilis. This adnate wing cannot be detached without injury.

THE ARTICULATE WING.

The articulate wing can be removed from the nut and can be replaced without injury. An ineffective form of this wing is seen in the Gerardianae and in P. pinea, where the blade is very short and the base has no effective grasp on the nut.

The base of the effective articulate wing contains hygroscopic tissue which acts with the hygroscopic tissue of the cone-scales. The dry conditions that open the cone and release the seeds cause the bifurcate base of the wing to grasp the nut more firmly.

This articulate wing is found in P. aristata and in all Hard Pines except P. pinea, longifolia and canariensis. The wing-blade is usually membranous throughout, but in some species there is a thickening of the base of the blade that meets the membranous apical part in an oblique line along which the wing is easily broken apart. This last condition attains in P. Coulteri and its associates a remarkable development.

Plate VI, fig. 72 shows the wingless seed of P. cembroides; fig. 73 represents the seed of P. flexilis, with a rudimentary wing; fig. 74 shows two seeds of P. strobus, intact and with the wing broken away; fig. 75 represents the articulate wing, whose bifurcate base when wet (fig. 76) tends to open and release the nut. When dry (fig. 77) the forks of the base, in the absence of the nut, close together and cross their tips; figs. 78, 79 show the peculiar reinforced articulate wing of P. Coulteri.

Such wide variation in so important an organ suggests generic difference. But here we are met by the association of the different forms in species evidently closely allied. The two Foxtail Pines are so similar in most characters that they have been considered, with good reason, to be specifically identical; yet the seed-wing of P. Balfouriana is adnate, that of P. aristata articulate. P. Ayacahuite produces not only the characteristic wing of the Strobi, adnate, long and effective, but also, in the northern variety, a seed with a rudimentary wing, the exact counterpart of the seed of P. flexilis. In both sections of the genus are found the effective adnate wing (Strobi and Longifoliae) and the inefficient articulate wing (Gerardianae and Pineae). A little examination of all forms of the seed will show that they blend gradually one into another.

The color of the wing is occasionally peculiar, as in the group Longifoliae. There is usually no constancy in this character, for the wing may be uniform in color or variously striated in seeds of the same species. The length and breadth of the seed-wing, being dependent on the varying sizes of the cone-scale, differ in the same cone. They are also inconstant in different cones of the same species, and of this inconstancy the seed of P. ayacahuite furnishes the most notable example.

THE WOOD. Plate VII.

With the exception of the medullary rays, a very small proportion of the whole, the wood of Pinus, as seen in cross-section (fig. 82), is a homogeneous tissue of wood-tracheids with interspersed resin-ducts. In tangential section the medullary rays appear in two forms, linear, without a resin-duct, and fusiform, with a central resin-duct. In radial section the cells of the linear rays are of two kinds, ray-tracheids, forming the upper and lower limits of the ray, characterized by small bordered pits, and ray-cells, between the tracheids, characterized by simple pits.

The walls of the ray-tracheids may be smooth or dentate; the pits of the ray-cells may be large or small. These conditions admit of four combinations, all of which appear in the medullary rays of Pinus, and of which a schematic representation is given in Plate VII. These combinations are

This, the simplest classification of Pine-wood, is not without exceptions. P. pinea of the Hard Pines resembles, in its wood-characters, P. Gerardiana and P. Bungeana of the Soft Pines. The dentate ray-tracheids of P. longifolia are not always obvious. The tracheids of P. luchuensis, according to Bergerstein (Wiesner Festschr. 112), have smooth walls. My specimen shows dentate tracheids. There is also evidence of transition from small to large pits (I. W. Bailey in Am. Nat. xliv. 292). Both large and small pits appear in my specimen of P. Merkusii.

Of other wood-characters, the presence or absence of tangential pits in the tracheids of the late wood establishes a distinction between Soft and Hard Pines. These pits, however, while always present in Soft Pines, are not always absent in Hard Pines. The single and multiple rows of resin-ducts in the wood of the first year may prove to be a reliable sectional distinction, but this character has not been sufficiently investigated to test its constancy. The wood-characters, therefore, however decisive they may be for establishing the phylogenetic relations of different genera, must be employed in the classification of the Pines with the same reservations that apply to external characters.

Ray-tracheids with dentate walls and ray-cells with large pits are peculiar to Pinus. Therefore the presence of these characters, alone or in combination, is sufficient evidence for the recognition of Pine-wood. But the combination of smooth tracheids with small pits (subsection Paracembra) Pinus shares with Picea, Larix and Pseudotsuga.

Among Hard Pines the size of the pits has a certain geographical significance. The large pits are found in all species of the Old World except P. halepensis and P. pinaster; the small pits in all species of the New World except P. resinosa and P. tropicalis. The Asiatic P. Merkusii with both large and small pits is not strictly an exception to this geographical distinction. The four exceptional species by this and by other characters unite the Hard Pines of the two hemispheres.

THE BARK.

Bark is the outer part of the cortex that has perished, having been cut off from nourishment by the thin hard plates of the bark-scales. In the late and early bark-formation is found a general but by no means an exact distinction between Soft and Hard Pines. In the Soft Pines the cortex remains alive for many years, adjusting itself by growth to the increasing thickness of the wood. The trunks of young trees remain smooth and without rifts. In the Hard Pines the bark-formation begins early and the trunks of young trees are covered with a scaly or rifted bark. The smooth upper trunk of older trees is invariable in Soft Pines, but in Hard Pines there are several exceptions to early bark-formation. These exceptions are easily recognized in the field, and the character is of decisive specific importance (glabra, halepensis, etc.).

Among species with early bark-formation are two forms of bark: 1, cumulative, sufficiently persistent to acquire thickness and the familiar dark gray and fuscous-brown shades of bark long exposed to the weather; 2, deciduous, constantly falling away in thin scales and exposing fresh red inner surfaces. The latter are commonly known as Red Pines, as distinguished from Black Pines with dark cumulative bark. Deciduous bark changes after some years to cumulative bark, and the upper trunk only of mature trees is red. Red Pines, although usually recognizable by their bark, are by no means constant in this character. Oecological or pathological influences may check the fall of the bark-scales, and then the distinction between the upper and lower parts of the trunk becomes lost.

SUMMARY

The various characters that have been considered in the previous pages may be classified under different heads, some of them applicable to the whole genus, others to larger or smaller groups of species.

GENERIC CHARACTERS

Several characters, quite distinct from those of other genera, are common to all the species.

1. The primary leaf—appearing as a scale or bract throughout the life of the tree.
2. The bud—its constant position at the nodes.
3. The internode—its three distinct divisions.
4. The secondary leaves—in cylindrical fascicles with a basal sheath.
5. The pistillate flower—its constant nodal position and its verticillate clusters.
6. The staminate flower—its constant basal position on the internode and its compact clusters.
7. The cone—its clearly defined annual growths.

Pinus is also peculiar in the dimorphism of shoots and leaves and in their constant interrelations with the diclinous flowers. Evolutionary processes develop features peculiar to Pinus alone (the oblique cone, etc.), but confined to a limited number of species.

SECTIONAL CHARACTERS

There are several characters that actually or potentially divide the genus into two distinct sections, popularly known as Soft and Hard Pines.

• 1. The fibro-vascular bundle of the leaf, single or double.
• 2. The base of the bract subtending the leaf-fascicle, non-decurrent or decurrent
• 3. The phyllotaxis of the cone, simple or complex.
• 4. The flower-bud, its less or greater development.

Some characters indicate the same distinction but are subject each to a few exceptions.

• 5. The fascicle-sheath, deciduous or persistent.
• 6. The walls of the ray-tracheids, smooth or dentate.
• 7. The connective of the pollen-sacs, large or small.
• 8. The formation of bark, late or early.

SUBSECTIONAL CHARACTERS

An exact subdivision of the Soft Pines is possible on the following characters.

1. The umbo of the cone-scales, terminal or dorsal.
2. The scales of the conelet, mutic or armed.
3. The pits of the ray-cells, large or small.

EVOLUTIONAL CHARACTERS

The progressive evolution of the fruit of Pinus, from a symmetrical cone of weak tissues, bearing a wingless seed, to an indurated oblique cone with an elaborate form of winged seed and an intermittent dissemination, appears among the species in various degrees of development as follows—

These different forms of the seed and, to some extent, of the cone, are available for segregating the species into groups of closely related members; while the gradual progression of the fruit, from a primitive to a highly specialized form of cone and method of dissemination, points to a veritable taxonomic evolution which is here utilized as the fundamental motive of the systematic classification of the species.

SPECIFIC CHARACTERS

All aspects of vegetative and reproductive organs may contribute toward a determination of species, but the importance of each character is often relative, being conclusive with one group of species, useless with another. Characters considered by earlier authors to be invariable with species, such as the dimensions of leaf or cone, the number of leaves in the fascicle, the position of the resin-ducts, the presence of pruinose branchlets, etc., prove to be inconstant in some species. In fact, as the botanical horizon enlarges, the varietal limits of the species broaden and many restrictions imposed by earlier systems are gradually disappearing.

Variation is the preliminary step toward the creation of species, which come into being with the elimination of intermediate forms. Variation in a species may be the result of its participation in the evolutionary processes culminating in the serotinous Pines, or it may result from the ability of the species to adapt itself to various environments by sympathetic modifications of growth, or it may arise from some peculiarity of the individual tree.

Evolutionary variation is associated with the gradual appearance of the persistent, the oblique and the serotinous cone, and of the multinodal spring-shoot. For these conditions appear in less or greater prevalence among the species of the genus.

Variation induced by environment finds familiar illustrations among the species that can survive at the limits of vegetation and can meet these inhospitable conditions by a radical change of all growing parts. Such variations are mainly of dimensions, but, with some species, the number of fascicle-leaves is affected and the shorter growing-season may modify the cone-tissues. In Mexico and Central America are found extremes of climate within small areas and easily within the range of dissemination from a single tree. The cause of the bewildering host of varietal forms, connecting widely contrasted extremes, seems to lie in the facile adaptability of those Pines, which are able to spread from the tropical base of a mountain to a less or greater distance toward its snow-capped summit.

The peculiarities of individual trees that induce abnormally short or long growths, the dwarf or other monstrous forms, the variegations in leaf-coloring, etc., etc., are not available for classification, for they may appear in any species, in fact in any genus of Conifers. These variations are artificially multiplied for commercial and decorative purposes. But inasmuch as they are repeated in all species and genera of the Coniferae that have been long under the observation of skillful gardeners, their significance has a broader scope than that imposed by the study of a single genus.