The Genus Pinus

Part 2 Classification of the Species 22

Sections, subsections and groups 25

Section Haploxylon 26

Subsection Cembra 26

Group Cembrae 26

Pinus Koraiensis, Cembra, Albicaulis 26, 27 VIII

Group Flexiles 28

Pinus Flexilis, Armandi 28, 30 IX

Group Strobi 30

Pinus Ayacahuite, Lambertiana 30, 32 X

Parviflora, Peuce, Excelsa 32, 34 XI

Monticola, Strobus 34, 36 XII

Subsection Paracembra 36

Group Cembroides 38

Pinus Cembroides, Pinceana, Nelsonii 38, 40 XIII

Group Gerardianae 40

Pinus Bungeana, Gerardiana 40, 42 XIV

Group Balfourianae 42

Pinus Balfouriana, Aristata 42, 44 XV

Section Diploxylon 44

Subsection Parapinaster 44

Group Leiophyllae 44

Pinus Leiophylla, Lumholtzii 44, 46 XVI

Group Longifoliae 46

Pinus Longifolia, Canariensis 46, 48 XVII

Group Pineae 48

Pinus Pinea 48 XVIII

Subsection Pinaster 50

Group Laricionea 51

Pinus Resinosa, Tropicalis 51, 52 XIX

Massoniana, Densiflora 52 XX

Sylvestris, Montana 54 XXI

Luchuensis, Thunbergii, Nigra 56, 58 XXII

Merkusii, Sinensis, Insularis 58, 60 XXIII

Group Australes 62

Pinus Pseudostrobus 62 XXIV

Montezumae 64 XXV

Ponderosa 66 XXVI

Teocote, Lawsonii 68 XXVII

Occidentalis, Palustris 70 XXVIII

Caribaea 70 XXIX

Taeda, Glabra, Echinata 72, 74 XXX

Group Insignes 76

Pinus Pringlei, Oocarpa 76, 78 XXXI

Halepensis, Pinaster 78, 80 XXXII

Virginiana, Clausa 80 XXXIII

Rigida, Serotina, Pungens 82, 84 XXXIV

Banksiana, Contorta 84 XXXV

Greggii, Patula 86 XXXVI

Muricata, Attenuata, Radiata 86, 88 XXXVII

Group Macrocarpae 90

Pinus Torreyana, Sabiniana 90 XXXVIII

Coulteri 93 XXXIX

INDEX 94

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 associates. This arrangement involves no radical change in existing systems. The new associations 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 identity. The remarkable variation in the species may be attributed partly to this adaptability, partly to a participation, more or less pronounced, in the evolutionary processes that culminate in the serotinous Pines.

PART I. Characters of the Genus

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 assume 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 quantity, etc., assist in the diagnosis of species. Occasionally a peculiar bud, like that of P. palustris, 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. palustris and caribaea. Both produce multinodal shoots, but the former so rarely that it should be classed 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 associated 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.

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