The Genus Pinus.

by George Russell Shaw.

INTRODUCTION

This discussion of the characters of Pinus is an attempt to determine their taxonomic significance and their utility for determining the limits of the species. A systematic arrangement follows, based on the evolution of the cone and seed from the comparatively primitive conditions that appear in Pinus cembra to the specialized cone and peculiar dissemination of Pinus radiata and its a.s.sociates. This arrangement involves no radical change in existing systems. The new a.s.sociations in which some of the species appear are the natural result of another point of view.

Experience with Mexican species has led me to believe that a Pine can adapt itself to various climatic conditions and can modify its growth in response to them. Variations in dimensions of leaf or cone, the number of leaves in the fascicle, the presence of pruinose branchlets, etc., which have been thought to imply specific distinctions, are often the evidence of facile adaptability. In fact such variations, in correlation with climatic variation, may argue, not for specific distinction, but for specific ident.i.ty. The remarkable variation in the species may be attributed partly to this adaptability, partly to a partic.i.p.ation, more or less p.r.o.nounced, in the evolutionary processes that culminate in the serotinous Pines.

PART I

CHARACTERS OF THE GENUS

THE COTYLEDON. Plate I, figs. 1-3.

The upper half of the embryo in Pinus is a cylindrical fascicle of 4 to 15 cotyledons (fig. 1). The cross-section of a cotyledon is, therefore, a triangle whose angles vary with the number composing the fascicle.

Sections from fascicles of 10 and of 5 cotyledons are shown in figs. 2 and 3. Apart from this difference cotyledons are much alike. Their number varies and is indeterminate for all species, while any given number is common to so many species that the character is of no value.

THE PRIMARY LEAF. Plate I, figs. 4-6.

Primary leaves follow the cotyledons immediately (fig. 4) and a.s.sume the usual functions of foliage for a limited period, varying from one to three years, secondary fascicles appearing here and there in their axils. With the permanent appearance of the secondary leaves the green primaries disappear and their place is taken by bud-scales, which in the spring and summer persist as scarious bracts, each subtending a fascicle of secondary leaves. At this stage the bracts present two important distinctions.

1. The bract-base is non-decurrent, like the leaf-base of Abies fig. 5.

2. The bract-base is decurrent, like the leaf-base of Picea fig. 6.

The two sections of the genus, Haploxylon and Diploxylon, established by Koehne on the single and double fibro-vascular bundle of the leaf, are even more accurately characterized by these two forms of bract-insertion. The difference between them, however, is most obvious on long branchlets with wide intervals between the leaf-fascicles.

The bracts of spring-shoots are the scarious bud-scales of the previous winter; but the bracts of summer-shoots have the form and green color of the primary leaf.

THE BUD. Plate I, figs. 7-11.

The winter-bud is an aggregate of minute buds, each concealed in the axil of a primary leaf converted into a scarious, more or less fimbriate, bud-scale. Buds from which normal growth develops appear only at the nodes of the branches. On uninodal branchlets they form an apical group consisting of a terminal bud with a whorl of subterminal buds about its base. On multinodal branchlets the inner nodes bear lateral buds which may be latent.

Fig. 7 represents a magnified bud of P. resinosa, first immersed in alcohol to dissolve the resin, then deprived of its scales. This bud contains both fascicle-buds, destined for secondary leaves, and larger paler buds at its base. These last are incipient staminate flowers, sufficiently developed for recognition. Such flower-bearing buds are characteristic of the Hard Pines in distinction from the Soft Pines whose staminate flowers cannot be identified in the bud.

The want of complete data leaves the invariability of this distinction in question, but with all species that I have examined, the flowers of Hard Pines are further advanced at the end of the summer. In the following year they open earlier than those of Soft Pines in the same locality. The staminate flowers of some Hard Pines (resinosa, sylvestris, etc.,) are not apparent without removing the bud-scales, but, with most Hard Pines, they form enlargements of the bud (fig. 9).

Invisible or latent buds are present at the nodes and at the apex of dwarf shoots. The former are the origin of the numerous shoots that cover the trunk and branches of P. rigida, leiophylla and a few other species (fig. 10). The latter develop into shoots in the centre of a leaf-fascicle (fig. 11) when the branchlet, bearing the fascicle, has been injured.

The size, color and form of buds, the presence of resin in quant.i.ty, etc., a.s.sist in the diagnosis of species. Occasionally a peculiar bud, like that of P. pal.u.s.tris, may be recognized at once.

THE BRANCHLET. Plate I, figs. 12-14.

The branchlet, as here understood, is the whole of a season's growth from a single bud, and may consist of a single internode (uninodal, fig.

12-a) or of two or more internodes (multinodal, fig. 13), each internode being defined by a leafless base and a terminal node of buds.

The spring-shoot is uninodal in all Soft Pines and in many Hard Pines, but, in P. taeda and its allies and in species with serotinous cones, it is more or less prevalently multinodal.

The uninodal spring-shoot may remain so throughout the growing season and become a uninodal branchlet. Or a summer-shoot may appear on vigorous branches of any species with the result of converting a uninodal spring-shoot into an imperfect multinodal branchlet. The summer-shoot may be recognized, during growth, by its green, not scarious bracts and, at the end of the season, by the imperfect growth of its wood and foliage (fig. 14).

The perfect multinodal branchlet is formed in the winter-bud (fig. 8-a) and the spring-shoot is multinodal. It is gradually evolved among the Hard Pines, where it may be absent, rare, frequent or prevalent, according to the species. In fact there is, in Pinus, an evolutionary tendency toward multinodal growth, with its beginnings in the summer-shoot and its culmination in the multinodal winter-bud, most prevalent among the serotinous Pines.

The multinodal shoot is never invariable in a species, but is rare, common or prevalent. This condition prevents its employment for grouping species. For Pines are not sharply divided into multinodal and uninodal species, and no exact segregation of them, based on this difference, is possible. In fact the character is unequally developed among closely related species, such as P. pal.u.s.tris and caribaea. Both produce multinodal shoots, but the former so rarely that it should be cla.s.sed as a uninodal species, while the latter is characteristically multinodal.

The multinodal spring-shoot, however, has a certain correlative value in its relation to other evolutionary processes that are obvious in the genus.

The length of the branchlet is much influenced by different soils and climates. In species able to adapt themselves to great changes, the length of the internode may vary from 50 cm. or more to 1 cm. or less.

In the latter case the branch is a series of very short leafless joints terminated by a crowded penicillate tuft of leaves (fig. 12-b). Such a growth may be seen on any species (ponderosa, albicaulis, resinosa, etc.) that can survive exposure and poor nourishment.

The presence of wax, as a bloom on the branchlet, is a.s.sociated with trees in arid localities, especially Mexico, where it is very common.

With several species the character is inconstant, apparently dependent on environment, and is a provision against too rapid transpiration.

The branchlet furnishes evidence of the section to which the species belongs, for the bract-bases persist after the bracts have fallen away.

The color of the branchlet, its l.u.s.tre, the presence of minute hairs, etc., are often suggestions for determining species.

[Ill.u.s.tration: PLATE I. PRIMARY LEAF, BUD AND BRANCHLET]

THE SECONDARY LEAF. Plate II.

Secondary leaves, the permanent foliage of Pines, are borne on dwarf-shoots in the axils of primary leaves. They form cylindrical fascicles, rarely monophyllous, prevalently of 2, 3 or 5 leaves, occasionally of 4, 6, 7, or 8 leaves. The scales of the fascicle-bud elongate into a basal sheath, deciduous (fig. 15) in all Soft Pines except P. Nelsonii, persistent (fig. 16) in all Hard Pines except P.

leiophylla and Lumholtzii. Inasmuch as these three species are easily recognized, the fascicle-sheath is useful for sectional distinctions.

EXTERNAL CHARACTERS.

The number of leaves in the fascicle is virtually constant in most species, the variations being too rare to be worthy of consideration.

With some species, however, heteromerous fascicles are normal. The influences that cause this variation are not always apparent (echinata, etc.), but with P. ponderosa, leiophylla, sinensis and others, the number of leaves in the fascicle is, in some degree, dependent on climatic conditions, the smaller number occurring in colder regions. In Mexico, for example, where snow-capped mountains lie on subtropical table-lands and extremes of temperature are in juxtaposition, the conditions are favorable for the production of species with heteromerous fascicles, and the number of leaves in the fascicle possesses often climatic rather than specific significance.

Among conifers, the leaf of Pinus attains extraordinary length with great variation, from 5 cm. or less to 50 cm. or more, the maximum for each species being usually much more than twice the minimum. Climate is the predominating influence; for the shortest leaves occur on alpine and boreal species, the longest leaves on species in or near the tropics.

The length of the leaf is complicated by the peculiarities of individual trees and by pathological influences; as a general rule, however, the length of leaves is less or greater according to unfavorable or favorable conditions of temperature, moisture, soil and exposure.

Therefore the dimensions of the leaf may be misleading. It can be said, however, that certain species always produce short leaves, others leaves of medium length, and others very long leaves.

Persistence of the leaf varies with the species and with the individual tree. But it is noteworthy that the longest persistence is a.s.sociated with short leaves (Balfouriana, albicaulis, montana, etc.).

INTERNAL CHARACTERS.

Since the leaf-fascicle is cylindrical, the cross-section of a leaf is a sector, its proportional part, of a circle. Theoretically the leaf, in section, should indicate the number of leaves composing its fascicle.

This is absolutely true for fascicles of two leaves only. No fascicle of five leaves, that I have examined, is equally apportioned among its five members. It may be divided in various ways, one of which is shown in fig. 18, where the leaf (a) might be mistaken for one of a fascicle of 3, and the leaf (b) for one of a fascicle of 6. Therefore if absolute certainty is required, a fascicle of triquetral leaves is best determined by actual count.