A handbook of systematic botany

The following characters should be added to those already given on page 2:—

The Gymnosperms comprise only trees or shrubs. The flowers are always unisexual and destitute of perianth (except Gnetaceæ); the female plant of Cycas is the only one which has no flower. The MALE FLOWERS are constructed on the same type as the cones of the Horsetails and Club-Mosses, and are most frequently long shoots (Figs. 243, 258, 260 A, 267 J) bearing a number of spiral or verticillate stamens. The FEMALE FLOWERS are of a more varied structure (see the orders). The OVULE is orthotropous (except Podocarpus which is anatropous) and projects from the carpel uprightly, inverted, or horizontally; it has usually only one integument (compare however Taxaceæ) which proceeds from the upper part of the nucellus, so that the embryo-sac in part is placed below the integuments (Figs. 251, 264). The drop of mucilage which catches the pollen-grains dries up and draws the pollen-grain through the micropyle to a space just above the nucellus—the pollen-chamber—in which the germination of the pollen-grain commences.

In each seed, only one of the many embryos which are formed proceeds to its full development. The seed is always endospermous, and the embryo has one, two, or a whorl of several cotyledons. A vigorous primary root is developed on germination. The vascular bundles in the stem are arranged in a ring, and increase in thickness takes place by a closed cambium-ring which forms bast (phlœem) externally, and wood (xylem) internally with distinct annual rings, as in the Dicotyledons. Only certain of the Cycadeæ deviate from this arrangement. The secondary wood is very uniform, as it is almost exclusively formed of tracheides with bordered pits, but true vessels are wanting; this also indicates a relationship with the Pteridophyta (see page 202).

The Gymnosperms are biologically lower than the Angiosperms; they are wind-fertilised, and without extra floral-nectaries.

This Division embraces three classes: Cycadeæ, Coniferæ, And Gneteæ. It is no doubt monophyletic, and has taken its origin from heterosporous Ferns, now extinct, most nearly related to the Ophioglossaceæ and Marattiaceæ. The Cycadeæ appear to be the oldest class. The Coniferæ are related to these through Ginkgo. The Gnetaceæ are more isolated. The Division is not continued into the higher Flowering-plants; it has evidently attained its highest development, and is now in a retrograde condition. The similarity which has often been pointed out between certain Coniferæ and Lycopodinæ is only in analogous resemblances, and does not entitle one to suppose that there is a nearer relationship, or that the former take their origin from the latter.

Class 1. Cycadeæ.

The stem is very rarely ramified. The leaves are large, pinnate, and arranged spirally. The flowers are diœcious, without perianth.

There is only one order, the Cycadaceæ.—In habit they resemble the Ferns, especially the Tree-Ferns (compare Figs. 207 and 256). The stem is tubercular (Fig. 258), or cylindrical (Fig. 256), but not very tall (as much as about 12 metres), and very rarely ramified. [In Ceylon, unbranched specimens of Cycas are rarely met with in the wild state. The stems of C. circinalis occasionally branch in greenhouses.]

The LEAVES are arranged spirally, and so closely together that no free stem-surface is left between them, and have only a slight sheath (which is not amplexicaul, as in the Palms). They are compound (most frequently pinnate; in Bowenia, bipinnate); in some genera the leaves are rolled up in various ways, resembling the vernation in Ferns (Fig. 257); they are leathery and perennial. In some, stipules are present, as in the Marattiaceæ. Groups of scale-leaves alternate in the majority with groups of foliage-leaves.

The FLOWERS are without perianth. The MALE FLOWER is sometimes an enormous collection of stamens (Fig. 258), which are flat in some (Cycas, Fig. 241), shield-like in others (Zamia, Ceratozamia) like the sporophylls in Horsetail (Fig. 259); but in all, the pollen-sacs are situated in large and varying numbers on the back of the stamens, and arranged in groups of 2–5, like the sporangia in the sori of the Ferns (Fig. 241 b, c). Female flowers are wanting in Cycas, because the carpels do not terminate the apical growth of the stem. After a group of foliage-and of scale-leaves, a group of carpels is developed, which are pinnate and resemble the foliage-leaves, bearing on their edges a number of ovules (most frequently 5–6) (Figs. 245, 256); the same stem produces successively scale-leaves, foliage-leaves, and carpels. The differentiation is not much more advanced than in certain Ferns (Struthiopteris, Blechnum), where barren and fertile leaves of different form regularly alternate. The other genera have female flowers; the carpels are shield-like in Zamia and Ceratozamia (Fig. 246), and collected into cone-like flowers, which terminate the growth of the stem (Fig. 259). The number of ovules in these instances is two to each carpel.

The SEEDS are large (most frequently 2–6 centimetres long) and plum-like; the external layer of the testa is fleshy, while the internal one is hard and horny. There are two systems of vascular bundles in the testa, one outside, the other inside the stone. The embryo is straight, attached to the end of the suspensor, which is often long, filamentous, and rolled up; it has one or two cotyledons.

Class 2. Coniferæ (Pine-trees).

The stem branches freely. The leaves are entire, relatively small, linear or reduced to scales. The flowers are without perianth. The ovules naked. It is seldom that the female flower is reduced to only one carpel.

Whilst the Cycadeæ principally resemble the Ferns, the Conifers partly resemble the Lycopods, and partly the Equisetums—the former especially in the needle- or scale-like, leathery, simple, and often perennial leaves (“evergreen plants”), which never possess stipules (Figs. 263, 270, 272). Ginkgo deviates from this, being no doubt the oldest, and the Conifer which stands nearest to the Cycadeæ (Fig. 260). The resemblance to the Equisetums is especially owing to the fact that the stem ramifies abundantly, and often very regularly, forming a pyramid with verticillate branches. In addition to the foliage-leaves, scale-leaves (bud-scales) are present in the majority of species.

The FLOWERS are monœcious or more rarely diœcious. Perianth is wanting. The stamens of the catkin-like male flowers (Fig. 267, J) are of different forms, but as a rule more or less shield-like. As in the Cycadeæ, the pollen-sacs are in all cases situated on the underside. There are, as a rule, two pollen-sacs (the Abietaceæ, Fig. 267), or 3–5, (the Cupressaceæ and Taxaceæ, Fig. 243); a few have more, e.g. Araucaria (Fig. 242); they dehisce by clefts.

If, in commencing our consideration of the female flower, we begin with that of Ginkgo, we shall observe in the corner of a scale- or foliage-leaf a small flower, which consists of two carpels, each bearing one ovule, and reduced almost to the ovule itself (Fig. 260 C, D). The flower in Podocarpus is still further reduced, viz. to a single carpel with one ovule, which is anatropous and has two integuments. This ovule is situated in the axil of a cover-scale (c, in Fig. 262 D), and several female flowers of this description are collected in a small cone, the stalk and bracts of which become fleshy (Fig. 262 C). The external integument also becomes fleshy (an aril). Dacrydium, which is clearly related to Podocarpus, has an external integument which developes more independently as a fleshy aril (Fig. 262 B, B’). Microcachrys also is clearly allied to these: the bracts are more fleshy, and the ovule (i.e. the female flower) is protruded beyond the bract (Fig. 262 A, A’). Taxus stands in a more isolated position: a flower which has been reduced to an ovule is situated, in this instance, on the apex of a secondary branch which is studded with floral-leaves (Figs. 263, 264); an external integument is developed on all sides and surrounds the seed as a scarlet aril. According to this conception the aril corresponds to an external integument, and the Taxoideæ thus possess a partly dichlamydeous ovule. Only Ginkgo and Cephalotaxus appear to deviate from this, as in these there is only one integument (unless the small outgrowth indicated by ar, in Fig. 260 D, really is a rudimentary, external integument); in Cycadeæ, to which Ginkgo is most closely related, there is likewise only one integument. But in these genera the testa is differentiated into two layers, and the seed resembles a drupe; like the Cycadeæ there is an external fleshy covering and an internal hard one, and these two layers may probably be considered homologous with the two integuments. This theory is also borne out by the arrangement of the vascular bundles in Cephalotaxus and Podocarpus, which present the xylem in the fleshy external layer to the outside of the testa, which is therefore the upper side of the integument (Celakovsky).

The coalescence of the integuments into one is only slight in Torreya, more pronounced in Podocarpus and strongest in Cephalotaxus and Ginkgo. Celakovsky terms these ovules “holochlamydeous.”

If we pass from these to the order Pinoideæ, we find the female flowers collected into catkin-like cones, which have been considered from various points of view to be sometimes single flowers, at other times compound inflorescences. The structure in Abietaceæ is as follows: a number of spirally arranged, scale-like leaves, cover-scales (Figs. 267, 268), are situated on a long axis. In the axil of each cover-scale a larger leaf-like projection, the ovuliferous scale, is borne, which turns the upper side towards its cover-scale (which is shown by the fact that the wood of its vascular bundles is turned downwards and towards the wood in the bundles of the cover-scale: Fig. 269). Two ovules, with micropyles turned towards the central axis, and with apparently only one integument (Fig. 268), are situated on the dorsal side of each ovuliferous scale, i.e. the side turned away from the cover-scale. The ovuliferous scales grow after fertilisation, into the woody or leathery “cone-scales,” which are usually much larger than the cover-scales. This ovuliferous scale with its axis may, according to Celakovsky, be considered as a dwarf-branch which is situated in the axil of the cover-scale, and bears two ovules (in the same way as in Ginkgo, one long-stalked flower, reduced to two ovules, is situated in the axil of a leaf), and in this case the external integument of the ovules is expanded into leaf-like bodies, which have united to form one “symphyllodium” (ovuliferous scale) which is inverted so that its dorsal side is turned upwards and bears the nucellus and the other integument (“hemichlamydeous” ovules). The carpel itself is therefore in this instance extremely reduced. The keel, or (in Pinus) “mucro” (Fig. 268 B), which is found in several genera, represents then a third carpel, which is sterile. In the other orders of the Pinoideæ the cover-scales and ovuliferous scales grow more and more together and finally form one structure, which also is termed a “cone-scale,” although from its development it cannot be homologous with the cone-scales of the Abietaceæ. This connation is least in the Taxodiaceæ and Araucariaceæ and may be traced on the upper surface of the “cone-scale” by the presence of a stronger or slighter ridge or pad, the free portion of the ovuliferous scale (Figs. 256, 266, 269). It is most strongly pronounced in the Cupressaceæ, in which the two scales form one single structure, the cone-scale (Fig. 274). The vascular bundles in the under portion corresponding to the cover-scale, have the xylem towards the upper side as usual in leaves, whilst the bundles present in the upper side of the cone-scale, which thus represents the ovuliferous scale, turn their xylem downwards. The hemichlamydeous ovules are then situated on the upper side of this cone-scale. According to this theory the Cupressaceæ appear to be the youngest type, a view which corresponds with their vegetative structure. If there is only one ovule in these orders as in Agathis (Fig. 265) and Araucaria, then the flower is reduced to a single carpel and one ovule, as in the case of Dacrydium and Microcachrys. If two or more ovules are present, then the same number of carpels may be supposed to exist, the external integuments of their ovules being developed into leaf-like structures which collaterally coalesce to form a “symphyllodium,” or are suppressed.

According to this theory, which is based on the researches of Celakovsky, the female flowers of the Coniferæ may be classed thus:—

1. In all cases situated in the axil of a bract and collected into cones, with numerous flowers or with few or one flower. In Ginkgo only, are they situated in the axil of foliage- or scale-leaves.

2. It is only in Taxus that bracteoles are present.

3. They are formed only from rudimentary carpels, in which the stem takes no part.

4. The number of carpels in each flower varies from one to many, most frequently three, of which the central one remains sterile.

5. Each carpel bears only one ovule. The flower which is formed of only one carpel appears to consist of only one ovule.

6. The ovule has in Taxaceæ either a double integument (Podocarpeæ, Taxeæ), of which the external is the “aril,” or, as in the Cycadeæ, a single one, which is homologous with the two united together.

7. The external integument in the Pinoideæ is expanded to form a leaf-like structure—the ovuliferous scale—and bears on its dorsal side the ovules, which are thus only provided with one, and that the inner, integument.

Pollination is accomplished by means of the wind. At the period of pollination the leaves are always so widely separated from one another, that the ovules can catch the pollen-grains carried to them by the wind; this is often effected by the mucilaginous drops which appear at the micropyle, and by the evaporation of which the pollen-grains are brought in contact with the nucellus. The entire cone grows considerably as soon as fertilisation has taken place, and the cone-scales in Pinoideæ close together so that the seeds while maturing are enclosed, and it is not until the seeds are ready for distribution that the cone-scales again become separated. In the Pinoideæ, the fully developed ovuliferous scales are hard and woody; and in this condition the collection of female flowers is termed a cone. In the Taxoideæ, true cones are the exception. 2–15 cotyledons are present, arranged in a whorl.

The characteristic feature of this class is the abundance of resin, which is to be found in isolated cells (especially in the cortex), partly in intercellular glands or passages (both in the cortex and wood). Taxus is the only genus which has no resin.

This class may be divided into two families:—

1. Taxoideæ. The ovules have either one integument, the external part of which is fleshy, and the internal hard and stone-like; or two integuments, of which the external is the fleshy and coloured “aril.” “Ovuliferous scales” are wanting. The cones are never woody, but are generally succulent, the bracts become fleshy, or cones usually are not developed. The seeds project more or less freely beyond the bracts.

2. Pinoideæ. The ovules have two integuments, the external one of which is leaf-like and becomes developed as the “ovuliferous scale”; if there are several of these in each flower they unite and form a “symphyllodium.” This may remain free or unite with the bract. The cones are most frequently woody, rarely succulent. The seeds are hidden among the cone-scales.

Family 1. Taxoideæ.

This family, considered to be most nearly related to the Cycadeæ, also made its appearance at a very early period. There is only one order.

Order. Taxaceæ. The characters have been given above.

A. Cephalotaxeæ is the oldest group, presumably the connecting link between the Cycadeæ and the other Coniferæ. The flower consists normally of two ovules. Aril wanting. One integument. Seeds drupaceous.—The flowers in Ginkgo biloba (Salisburia) are situated in the axil of foliage- or scale-leaves. The stamens bear only two pollen-sacs (Fig. 260 A). The female flower has two ovules, placed together at the end of a long, bare axis (Fig. 260 C). Round the base of the ovule a small collar (ar, in Fig. 260 D) is found, which may probably be considered homologous with the collar-like outgrowth which surrounds the base of the Cycas-ovule. The seed resembles a Plum, and has a fleshy external coat, surrounding a hard internal layer. The embryo is developed after the seed has fallen off. The Ginkgo-tree has long-stalked, fan-shaped leaves, more or less indented, with dichotomous veins resembling certain Ferns—the Adiantums. It is a native of East Asia, and the only surviving species of a genus which in earlier times was very rich in species, and distributed over the entire Northern Hemisphere. Cephalotaxus (Eastern Asia) is related to it.

B. Podocarpeæ. The female flower is reduced to one ovule, placed in the axil of a bract, or a little forward upon it. The ovule has an aril (2 integuments).—Phyllocladus (Fig. 261), from New Zealand and Tasmania, has obtained its name from its flat, leaf-like branches, the leaves proper being scale-like (f). The ovules stand erect in the axil of the scale-like leaves (c), and several are collected at the end of short branches.—Microcachrys tetragona (Tasmania) has a small female catkin with several spirally-placed, fleshy bracts, at the end of which the inward and downward turned ovule is attached (Fig. 262 A, A’). The ripe cones are red, succulent, and resemble Strawberries.—In Dacrydium (Tasmania, New Zealand, Malaysia) the female cone has most frequently only 1–2 (–6) bracts, which resemble the vegetative leaves; they have also a fleshy aril (Fig. 262 B, B’).—Podocarpus (40 species, East Asia, S. Temp.); the bracts of the female flowers become fleshy, and unite together; only 1 or 2 are of use in supporting the flowers. The ovules project high above the apex of the bract, and are anatropous, the micropyle being turned downwards (Fig. 262 C, D). An aril commences to develope in the flowering period as an external coating, and later on it becomes fleshy and coloured.

C. Taxeæ. The female flower is reduced to one ovule, which is situated terminally on an axis which bears 2–3 pairs of opposite, scale-like bracteoles; on this account the Taxeæ form a very isolated group among the Coniferæ.—Taxus (T. baccata, the Yew-tree). Diœcious. The female flower consists of only one ovule, placed at the end of a short secondary branch (Fig. 264), which is studded with scale-like leaves. The aril when ripe is thick, fleshy, and scarlet (sometimes yellow), and only loosely envelopes the seed (Fig. 263). The leaves are scattered, flat, linear, and pointed (Fig. 263, 264). The short male flowers have 5–8 pollen-sacs, pendent from the stamens, and are surrounded at their bases by scale-like bracteoles (Fig. 243). Torreya (4 species, N. America and Japan) is closely allied to Taxus. The aril ultimately fuses with the woody inner integument, and hence the ovule becomes drupaceous, as in Cephalotaxaceæ.

Family 2. Pinoideæ.

The four orders differ from one another partly in the arrangement of the leaves (Cupressaceæ have opposite or verticillately placed leaves, flowers, and inflorescences; in the others they are placed spirally), but chiefly in the greater or less degree of union which takes place between the female flower (the leaf-like “symphyllodium”) and its supporting cover-scale, and in the position of the ovules (the micropyle being turned upwards or downwards). The “cone-scales” in Abietaceæ are formed by “symphyllodia” alone, in the others by their union with the cover-scale.

Order 1. Araucariaceæ. This order most frequently has solitary ovules, turned downwards and attached to the centre of the cone-scales. In Agathis (Dammara) the arrangement is the most simple, a winged seed (Fig. 265), which hangs freely downwards, being borne in the centre of the undivided cone-scale. In Araucaria, the stamens with the free, pendulous pollen-sacs have been represented in Fig. 242; the ovuliferous scale is united for nearly its whole length with the bract, and projects from its apex in the shape of a sheath-like, dentate scale, resembling the ligule in Isoëtes, and may therefore be termed a “ligule.” Araucaria (S. America, Australia) has often rather broad leaves (A. brasiliensis). The ovuliferous scale in Cunninghamia is more distinct, and stretches transversely over the entire cover-scale; it bears three inverted ovules (Fig. 266) (Eastern Asia).

Order 2. Abietaceæ (Pine and Fir Trees). The leaves are spirally arranged and needle-like. The flowers are monœcious. The male flowers are long, and catkin-like, with numerous stamens, each bearing two oblong pollen-sacs. The pollen-grains are most frequently tri-lobed, having two bladder-like appendages, formed as outgrowths of the exospore, to assist in their distribution by the wind (Fig. 267 N). The bracts are arranged spirally. The union between the bract and the ovuliferous scale, which is found in the preceding order, is not in this instance so complete; these scales make their appearance as two free parts, and are attached only at their bases (Fig. 268); the lower portion, that is the cover-scale, in most instances remains quite small (Fir, Red Pine, and others), it is only in the “Noble Pine” (Abies) and Pseudotsuga douglasii, that it attains a greater length than the ovuliferous scale (Fig. 267, B-G). On the other hand the upper part, the ovuliferous scale (the vascular bundles of which have the bast turned upwards), grows strongly and elongates, especially after fertilisation, becoming woody or leathery; it is commonly termed the “cone-scale,” but is in reality only homologous with a part of the “cone-scale” in the other order of Pinoideæ. On the side of the ovuliferous scale, turned towards the axis, are situated two ovules with micropyles directed inwards. The seeds are most frequently provided with a false wing (a tissue-like part of the surface of the ovuliferous scale). Cotyledons, more than 2, verticillate. Fertilisation does not take place until some time after pollination. In Pinus, for instance, the pollen-tube only penetrates the nucellus for a short distance during the year of pollination, and then ceases its further growth, fertilisation not taking place until after the middle of the next year; whilst the seeds ripen about a year and a half after pollination. In the Larch and others, the seeds are mature in the autumn succeeding pollination.

Abies (Fir). The leaves are often (e.g. Ab. pectinata) displaced into 2 rows, flat and indented at the apex, with 2 white (wax-covered) lines on the under surface, in which the stomata are situated. The leaf-scars are nearly circular and do not project. The cones are erect. The cover-scales and the ovuliferous scales separate from the axis, to which they remain attached in other genera.—Tsuga has leaves like Abies, but by the slightly projecting leaf-scars, and cones with persistent scales, it forms the transition to Picea.—Pseudotsuga has leaves similar to those of Abies and persistent carpels as in Picea, but the cover-scales grow as in Abies and project beyond the ovuliferous scales (P. douglasii, Fig. 267). These two genera are considered as sub-genera of Abies.—Picea. The leaves project on all sides, square and pointed; the leaf-scars are rhombic, on projecting leaf-cushions. The cones are pendulous. The cover-scales are much shorter than the leathery, persisting ovuliferous scales.—The genus Larix (Larch) differs from all the others in having deciduous leaves (the three preceding have leaves which persist for eleven to twelve years). It has long-branches with linear foliage-leaves and short, thick, perennial dwarf-branches, which each year form a new rosette of foliage-leaves, similar to those on the long-branches. The male flowers and the erect cones resemble those of Picea, and are borne on dwarf-branches.—Cedrus (Cedar) resembles Larix to some extent, but has persistent leaves (C. libani, C. deodara).—Pinus (Pine) has long-branches and dwarf-branches. The leaves of the long-branches are scale-like and not green; the dwarf-branches have very limited growth, and persist for three years; they arise in the axils of the scales borne on the long-branches of the self-same year, and each bears 2–5 foliage-leaves, they are also surrounded at the base by a number of membranous bud-scales. The cone-scales have a thick, rhomboid extremity (the “shield”).

Order 3. Taxodiaceæ. The vegetative leaves and cone-scales are arranged spirally. The ovules (2–9) are situated either at the base of the ovuliferous scales, in which case they are erect; or at their centre, when they are generally more or less inverted. The ovuliferous scale is more or less united with the cover-scale, and projects beyond the surface of the cone-scale, like a comb (Fig. 269). The vascular bundles, which extend into the cover-scale, have the usual leaf-arrangement, viz. the wood placed above the bast; while those bundles which enter the ovuliferous scale have this arrangement of the bundles reversed.