Fig. 466.—Ruta. Longitudinal section of flower.
Fig. 467.—Ruta. Floral diagram.
4. Ruteæ. Ruta (Figs. 465–467) graveolens is an herbaceous, glaucous, strongly smelling plant with bipinnate leaves and yellow flowers; the terminal flower is 5-merous, the others 4-merous (S. Eur.).—Dictamnus; zygomorphic flower. The individual carpels of the fruit separate from each other, and dehisce like follicles, upon which the internal layer is detached elastically and springs out, carrying the seeds with it. Several species are ornamental plants.
5. Cusparieæ. American. Flowers often zygomorphic with gamopetalous corolla; stamens 5.—Ticorea; Galipea (G. officinalis; S. Am.; “Cortex angosturæ”); Cusparia; Almeidea.
B. The ovary is entire or only slightly grooved; the style is terminal, undivided. The fruit is most frequently a drupe or berry.
6. Toddalieæ. Ptelea; winged fruit. The buds are enclosed in the leaf-sheath. Skimmia; Phellodendron.
Figs. 468–470.—Citrus vulgaris.
Fig. 468.—Branch with compound leaves.
Fig. 469.—Transverse section of fruit.
Fig. 470.—Flowers (after the removal of the petals).
7. Aurantieæ, Orange Group. Fruit a berry with a leathery external layer.—The most typical flower is found for example in Limonia: S5, P5, A5 + 5, G5 (2–5).—Citrus has 4–5–8-merous flowers, a gamosepalous, dentate calyx, free petals, one whorl of stamens which are split irregularly into several bundles (Fig. 470). The fruit is a multilocular berry provided with a thick, tough, outer layer. The juicy pulp, which fills up the loculi and envelopes the seeds, is formed from many large-celled, juicy hair-structures which arise on the inner side of the walls of the loculi and by degrees entirely fill them up; the dissepiments remain thin, and form the partitions so easily separating from each other (Fig. 469). The seeds in many instances are remarkable for containing several embryos. The blade of the leaf is separated from the frequently winged stalk by a node (and hence is a compound leaf with only the terminal leaflet developed?) (Fig. 468); in other genera, as Triphasia, there is a fully developed trifoliate leaf. Thorns are frequently developed.—The species of this genus, which is a native of the warmer parts of S. E. Asia, are very hard to separate. The differences are found in the forms of the fruit, the leaves and the leaf-stalks, and in the number of stamens. Citrus medica, “Cedrat” (Ind.); C. limonum, “Citron,” “Lemon” (introduced into Italy in the 3rd to 4th century). Officinal: the fruits and essential oil of Lemon. C. aurantium from E. Asia, the Orange (introduced into Italy in the 14th century). C. vulgaris (Fig. 468), Bitter Orange (introduced into Europe at the time of the Crusades); the unripe Bitter Oranges, and peel of the Bitter Orange is officinal; it is from the flowers of this species especially that the essence of Neroli is made. C. limetta, C. bergamia, Bergamot; essence of Bergamot is officinal. C. decumana, Pomalo, a native of the Islands of the Pacific. About 780 species; chiefly tropical.
Order 4. Burseraceæ. Fruit a drupe; 1–5 stones. The bark, as well as the other parts, contain strong aromatic resins and balsams, and hence several species are used: the Myrrh tree, Commiphora (Balsamodendron) from Arabia and Africa; Officinal: Myrrha (Commiphora myrrha). Mecca-balsam from C. opobalsamum, Arabia; E. Africa. The Incense-tree (Boswellia) from the same parts of the globe and E. India. The incense of B. carteri is medicinal (Frankincense). The resin (Elemi) of Protium-species is officinal, and is used technically for varnish (S. Am.). Takamahaka-resin from Elaphrium (S. Am.) Protium (Icica); Amyris (1 carpel). 270 species; tropical.
Order 5. Zygophyllaceæ. The majority have opposite, pinnate leaves with stipules. Leaves without pellucid dots. The filaments have a scale on the inner side. The most important is Guaiacum officinale (West India), the wood (Lignum Vitæ) of which is very hard and heavy, this wood and Gum-guaiacum are officinal. Others have a peculiar repulsive smell and taste: the Creosote shrub (Larrea mexicana) and Zygophyllum simplex. Tribulus terrester is a common weed in S. Europe. Fagonia. Peganum harmala (South of Russia) yields a red dye.—110 species; especially in the Tropics; several species in sandy deserts. Nitraria.
Order 6. Simarubaceæ. This order is distinguished by the abundance of bitter substances which it contains (Quassine) especially in the bark and the wood. The wood of Quassia amara (Guiana, Antilles) is officinal; Picraena excelsa yields Jamaica Quassia; the bark of Simaruba, Simaba-species and others is used. Ailanthus glandulosa is a garden plant (pinnate leaves, winged fruit).—110 species. Tropical.
Order 7. Ochnaceæ. Flowers diplostemonous, 5-merous. The unilocular ovaries, which are individually free, project considerably into the air around the gynobasic style; 1 ovule in each loculus; the fruitlets are drupes. Shrubs; leaves alternate, with stipules. Ochna; Ouratea.—160 species; tropical; especially American.
Order 8. Anacardiaceæ. The ovary rarely contains more than 1 ovule, even though there be several loculi and several carpels; in Anacardium all the 10 stamens except one become suppressed. Resin passages.—Anacardium. The most peculiar feature is the development of the flower-stalk into a fleshy body about the form and size of a pear (A. occidentale from Trop. Am. and A. orientale from E. Ind.) which bears the kidney-shaped nut (the so-called “Cashew-nut”) on its apex. Mangifera indica (the Mango-tree, from E. Ind.) is cultivated in several tropical countries on account of its delicious drupe. Similarly, species of Spondias (S. dulcis, Pacific Islands, S. lutea). Several species of Rhus are ornamental shrubs in this country, for instance, R. typhina (N. Am.), R. cotinus (the Wig-tree, the barren flower-stalks of the panicles being feather-like and hairy); R. toxicodendron (Poisonous Sumach, from N. Am.) is poisonous. Chinese galls are produced by the sting of a leaf-louse (Aphis chinensis) on R. semialata (China), and Japanese wax is from the seeds of R. succedanea (Japan). Considerable quantities of Sumach (R. coriaria) are used in tanning and as a black dye. Officinal: the mastic resin of Pistacia lentiscus (the Mastic-tree, from the Mediterranean). The fruits of Pistacia vera (Syria) are edible; P. terebinthus and others yield turpentine.—450 species; tropical.
Order 9. Icacinaceæ. Flowers 4–5-merous; haplostemonous; receptacle convex or cup-like surrounding the gynœceum; in the (single) loculus of the ovary, 2 anatropous, pendulous ovules.—200 species; tropical.
The essential characters of this family are in the main the same as those of the Terebinthinæ and Gruinales. The flowers are hypogynous, perfect, with free petals, 5-merous (S5, P5, typically A5 + 5, all of which, however, are not generally developed; in our native orders there are only 7–8 stamens), and most frequently a 3-merous, 3-locular gynœceum (less frequently 2 or 5 carpels with as many loculi). In each loculus there are usually only 1–2 ovules. A deviation from the preceding families is the frequent zygomorphy of the flower, with, as a rule an oblique plane of symmetry (Fig. 471). When a disc is developed it is placed outside the stamens. The majority have no endosperm (Fig. 473).—The members of the family are nearly all trees.
The family is closely allied to the Terebinthinæ, but unlike this it never has aromatic properties, and differs also in the position of the nectary, in the flowers, which are often irregular with a reduction in the number of stamens, and in the ovule which is usually ascending with micropyle pointing downwards (the Terebinthinæ having the micropyle turned upwards), etc. It is also related to Frangulinæ, the Staphyleaceæ being the chief connecting link; but the Æsculinæ generally have compound leaves.
Order 1. Staphyleaceæ. Leaves opposite, often compound. Flowers regular, ☿, 5-merous in calyx and corolla, 5-stamened. The stamens are placed outside the nectary. Ovary syncarpous or 2–3-partite with free styles. The capsule is thin, bladder-like, 2–3-locular, opening at the apex, and has several very hard seeds with a shining testa without aril. Endosperm. Staphylea pinnata (S. Europe) and trifoliata (N. Am.) are cultivated in gardens; they have white flowers in pendulous, axillary racemes or panicles.—16 species.—Staphylea is found in the Tertiary of N. America.
Order 2. Melianthaceæ. Glaucous shrubs with scattered, pinnate leaves, and large stipules. Melianthus.—8 species; S. Africa.
Order 3. Sapindaceæ. Trees or shrubs, often climbing by tendrils (lianes with anomalous structure of the stem) and with compound leaves. The flowers, in most cases, are small, insignificant, and without scent, and in some polygamous and zygomorphic. S4–5, P4–5, A8 (less frequently 5–10) inside the nectary (disc); ovary generally 3-locular, with 1–2 ovules in each loculus (raphe ventral, micropyle turned downwards). Seed without endosperm, often with an aril. The embryo is often thick and curved (Fig. 473).
Figs. 471–473.—Æsculus hippocastanum.
Fig. 471.—Diagram of the flower and of a scorpioid cyme.
Fig. 472.—Flower in longitudinal section.
Fig. 473.—Seed in longitudinal section.
Æsculus (Horse-Chestnut). Trees with opposite, digitate, dentate leaves without stipules; the inflorescence is composed of unipared scorpioid cymes arranged in a pyramidal panicle (termed a thyrsus). The flowers are irregular, with an oblique plane of symmetry (through the 4th sepal, Fig. 471); there are 5 sepals, 5 free petals, of which the one lying between S3 and S5 is the smallest (see Fig. 471) and may be absent; stamens 7 (5 + 2), three being suppressed; gynœceum simple, 3-carpellary and 3-locular, with single style; of the two ovules one is ascending, the other descending (Fig. 472).—The fruit is a 3-valvate, sometimes spiny, capsule, with loculicidal dehiscence, the seed having a large hilum, a curved embryo without endosperm and united cotyledons (the radicle lies in a fold of the testa, Fig. 473). Æ. hippocastanum (Greece, Asia), introduced into cultivation about 300 years ago; the majority of the other species, e.g. Æ. pavia, etc., several of which are frequently cultivated in gardens, are from N. America. The flower of the Horse-Chestnut is adapted for bees, whose abdomen touches the anthers or style when visiting the flower. The flowers are protogynous.
The other Sapindaceæ have most frequently 4 sepals, 8 stamens, various fruits (septicidal capsule, nuts with or without wings, schizocarp), etc. Serjania, Cardiospermum, Sapindus, Koelreuteria, etc. (about 118 genera, 970 species). The seeds of Paullinia sorbilis contain caffeine, and are used as “Pasta guaranà,” in the North Western Brazils in the manufacture of a common drink. Nephelium (or Euphoria) litchi (with edible aril), and other species, from Asia.
Fig. 474.—Samara of Acer platanoides.
Order 4. Aceraceæ. This order is so closely allied to the Sapindaceæ, that some authorities have classed it with them. The main difference is in the regularity of the flowers, and the 2-merous gynœceum (in abnormal cases several carpels occur).—They are trees, and, like the Horse-Chestnuts, have opposite leaves without stipules; in Acer the leaves are palminerved, but imparipinnate in Negundo, a plant frequently cultivated in gardens. The flowers are often unisexual, polygamous (some species have ☿-, ♂-and ♀-flowers); sepals 5, petals 5 free, stamens 8 (that is, 5 + 5, but the two median ones are absent) inside a large disc. Fruit a samara (schizocarp) with 2 winged, nut-like fruitlets (Fig. 474). In each of the 2 loculi of the ovary are 2 ovules. Embryo curved, with thin, folded cotyledons. Endosperm absent.—The inflorescences are racemes with a more or less elongated main axis and terminal flower (which sometimes has 10 stamens); when the lateral branches are developed they are similar to the main axis. In some species both corolla and petal-stamens are suppressed. Acer is pollinated by insects, Negundo by the wind.—88 species; North Temperate zone. Acer in the Tertiary from the Oligocene. The following are native plants: Maple (Acer campestre), Sycamore (A. pseudoplatanus, doubtful native). Important as avenue trees and timber. Sugar is obtained from the spring sap of the Sugar Maple (N. Am.).
Order 5. Malpighiaceæ. A tropical (especially American) order closely related to the Aceraceæ, having often the same form of fruit (but 3-partite). Some species are lianes with anomalous stem-structure. Leaves opposite. The flowers are regular or obliquely zygomorphic (the plane of symmetry passing through sepal 3), with S5, P5, A5 + 5, G3; 1 pendulous ovule in each loculus. Important characteristics for identification are the numerous grandular structures on the sepals. Peculiar 2-spined hairs are found in some. Malpighia, Bunchosia, Galphimia, Tetrapterys, Heteropterys, etc.—About 600 species.
Order 6. Erythroxylaceæ. Sepals 5, petals 5 (with a ligular corona), 10 stamens in one bundle. Gynœceum 3-locular. Fruit a drupe. Tropical (especially American) trees and shrubs, the Coca-plant (Erythroxylon coca) being best known. Its leaves are considered by the inhabitants of Chile and Peru to be one of the indispensable necessaries of life; they are chewed, and possess intoxicating, exhilarating properties, and contain the alkaloid cocaine, which is frequently employed as a local anæsthetic.—103 species; chiefly in America.
Order 7. Vochysiaceæ. Trees; Trop. Am. 1 stamen.—140 species.
Order 8. Trigoniaceæ. Shrubs; Trop. Am.—30 species.
Order 9. Tremandraceæ. Polygalaceæ with regular flowers.—27 species. Australia.
Fig. 475.—Diagram of Polygala: d a gland in the posterior side of the flower; α and β the two caducous bracteoles.
Order 10. Polygalaceæ. Herbs or shrubs (some tropical species are lianes) with scattered (rarely opposite), simple and most frequently quite entire leaves, without stipules. The flowers are usually borne in terminal spikes or racemes, and are strongly zygomorphic (the plane of symmetry being median); they have 5 free sepals, the 2 lateral ones of which (4 and 5 in Figs. 475, 476) are very large, petaloid, and frequently project on each side like the “wings” of a Pea-flower; petals 5, of which the two lateral ones are wanting or rudimentary (dotted on Fig. 475), and the anterior “the keel” (Fig. 476 c) is large, hollow and boat-shaped, and frequently with a lobed or fimbriated edge (Fig. 476 A and B, c); stamens 8, the two median ones being absent, all united into a tube split along the back, which is also slightly united to the keel (the anthers, often 2 locular, open by pores, Fig. 476 B, st); the 2 median carpels form a bilocular ovary. 1 pendulous ovule in each loculus (Figs. 476 C, 475); capsule compressed with loculicidal dehiscence, rarely a nut. Polygala (Milk-wort).
470 species; distributed over the whole globe (none Arctic). Officinal: the root of P. senega, from N. Am. Some are used as ornamental plants.
Pollination. The flowers of Polygala are pollinated by insects (chiefly bees). The fimbriated processes of the anterior petal support the insect when it alights. The anthers lie on each side of the stigma in the pouch of the anterior petal; the apex of the style is spoon-shaped, and immediately behind it is a viscid stigmatic lobe. In reaching the honey the proboscis of the insect must come in contact with the pollen and the viscid stigma, by which it is rendered sticky; this ensures the pollen adhering to the proboscis and so being carried to other flowers.
Fig. 476.—Polygala amara. Parts of the flower (mag.) A Flower from side, 1-5 sepals: c keel; B flower from above spread out: st the 8 stamens; c fimbriated edge of “keel”; C ovary with style and stigma.
The plants belonging to this family, with very few exceptions, are trees or shrubs. The leaves are usually simple; stipules may be absent or present. The flowers in almost all the orders are small, green or whitish; they are always regular, 4-or 5-merous with 2–5 carpels, but never have more than 1 whorl of stamens, which in Rhamnaceæ and Ampelidaceæ are placed opposite the petals (typically 5 + 5 or 4 + 4 stamens, of which however either the external or internal whorl is always wanting); hypogynous or slightly perigynous, in Rhamnaceæ only strongly perigynous or epigynous; generally ☿; the calyx is inconspicuous; petals free or slightly united. Gynœceum simple; ovary generally multilocular; style short or entirely wanting. A disc is nearly always developed in the flower, but is found sometimes inside the staminal whorl, sometimes outside it or between the stamens. The ovules are apotropous (anatropous with dorsal or ventral raphe).
Order 1. Celastraceæ. Euonymus europæa (Spindle-tree) may be chosen as a type. It is a shrub with simple, opposite leaves and small caducous stipules. The small, greenish-yellow flowers, borne in regularly-branched dichasia, are regular, ☿, with 4 whorls, 4-(or 5-) merous in regular alternation. There is a thick disc upon which the polypetalous corolla (imbricate in the bud) and the stamens are borne, with a slightly perigynous insertion. The style is short and thick; the ovary has 2 erect ovules in each loculus. The fruit is a red, 4-valvate capsule with loculicidal dehiscence; the seeds are few in number, and have a large, red-yellow aril (developed from the micropyle). Embryo green, in a large, fleshy, white endosperm. The dingy yellow flowers are generally visited only by flies and ants for the sake of the honey secreted by the disc, and while they run about on the flowers they touch the anthers and stigmas, now with one part of the body, now with another. The flower is protandrous. The stigmas are not developed till several days after the opening of the anthers.—Celastrus, Cassine, Catha, etc.
38 genera; 300 species. Distributed over the entire globe, with the exception of the colder districts, and especially in the Tropics. Some are ornamental bushes (Euonymus japonica). The leaves of Catha edulis are used by the Arabs and Abyssinians in the same way as those of Coca by the Peruvians.
Order 2. Hippocrateaceæ. 150 species; tropical; chiefly lianes. S5, P5, A3, G3. Anthers extrorse.
Fig. 477.—Ilex aquifolium: magnified flower.
Order 3. Aquifoliaceæ (Hollies). The genus Ilex forms almost the entire order. (175 species out of 180; especially from S. Am.) They are shrubs or trees with scattered, leathery, simple leaves (in Ilex aquifolium, spiny) with very small stipules. The flowers are small, white, and borne in few-flowered inflorescences in the axils of the foliage-leaves; they are most frequently unisexual and diœcious. There are 4–5 sepals, petals, stamens and carpels in regular alternation; the calyx and corolla have their leaves slightly connate; stamens slightly adnate to the corolla; the ovary is generally almost spherical with a thick, sessile stigma (Fig. 477). This order deviates especially from Celastraceæ in the absence of the disc and in having only 1 (pendulous) ovule in each of the 4 loculi of the ovary, and in having a drupe with generally 4 stones. Embryo extremely small, at the apex of the large endosperm, with the radicle directed upwards.—3 genera.—I. aquifolium (Holly) principally on the coasts of European countries; from Norway to W. Denmark, and further westward. It is a common garden shrub with stiff, shining leaves and red fruits. Several South American species contain so much caffeine that they may be used as a beverage in the place of tea (I. paraguayensis, Paraguay tea, or Maté). The Holly does not contain caffeine.
Order 4. Ampelidaceæ (Vines). Shrubs with the stem swollen at the insertion of the petioles and climbing by tendrils borne opposite the leaves (Figs. 478, 479). The leaves are scattered (generally 1/2), stalked, stipulate, frequently palminerved and lobed, divided or compound. The small, greenish flowers are generally borne in paniculate inflorescences, whose position is the same as that of the tendrils (Fig. 478); they are hypogynous or slightly perigynous, ☿, with 4–5 sepals, petals, stamens (which, as in the Rhamneæ, are opposite the petals; Fig. 480 A, B) and 2 carpels. The calyx is very small, entire, or slightly dentate; corolla valvate, and in some falling off as a hood, since the individual parts remain united at the summit (Fig. 480 A). Between the stamens and gynœceum is situated an hypogynous disc, with 5 lobes alternating with the stamens (Fig. 480 A, B, E). In each loculus of the 2-locular ovary there are 2 erect ovules (E); the style is short or wanting. The fruit is a berry. The embryo is small and lies in a horny, sometimes slightly folded (ruminate) endosperm (Fig. 480 C, D).
Figs. 478–481.—Vitis vinifera.
Fig. 478.—Branch with bunch of grapes.
Fig. 479.—Diagram of the position of leaf and tendrils. The branch is divided into sections on the sympodial theory (the successive generations, I, II, III, IV, are alternately white and shaded); k buds.
Fig. 480.—A Flower throwing off the corolla; B flower after the removal of the corolla; C, D longitudinal and transverse section of seed; E longitudinal section of gynœceum; s calyx.
Fig. 481.—Diagram of branch and position of leaves; sl tendril; lt the main axis; ax stipules of the foliage-leaf shown below; g axillary-bud (the dwarf-branch); v its fore-leaf; l1 l2 its first two foliage-leaves with their stipules; lt1 long-branch in the axil of v (everything appertaining to this branch is entirely black); v1 the first leaf of this branch.
Vitis and Ampelopsis (5-merous flowers); Cissus (4-merous flower); Leea (without stipules, corolla gamopetalous). The inflorescence in Pterisanthes (E. Ind.) has a peculiar, flat, leaf-like axis, on the edges of which ♂-flowers are borne, and on the surface ♀-flowers.
The TENDRILS in Ampelidaceæ are modified branches, since they bear leaves and may be abnormally developed as branches with foliage-leaves, and finally the inflorescences are borne in the position of the tendrils, and tendrils are met with which are partly inflorescences. The explanation of the position of the tendril, namely, right opposite the foliage-leaf but without a subtending-leaf, has been much disputed. The relative positions are as follows: in Vitis vinifera the following two kinds of shoots and relative positions are found (the other species deviate in one or other particular), (a) Long-branches, which have 2 scale-leaves and a large number of foliage-leaves with a divergence of 1/2; opposite the lowest 3–5 foliage-leaves no tendrils are found, then follow: 2 foliage-leaves with tendrils, 1 without a tendril, 2 with and 1 without, etc., with great regularity. Buds are developed in the axils of the foliage-leaves (Fig. 479): these develop into (b) DWARF-BRANCHES, which commence with 1 laterally-placed scale-leaf (fore-leaf; Fig. 481 v) succeeded by several foliage-leaves with a divergence of 1/2 (in a plane at right angles to that of the mother-shoot), but the whole shoot is extremely small, and often dries up and drops off in the autumn, so that only the scale-leaf, v, with the bud (Fig. 481 lt1) in its axil remains. This bud in the following year developes into a new long-branch, and since its leaves lie in a plane at right angles to that of the dwarf-branch, their plane coincides with that of the long-branch from which it is developed (the grandmother axis).—The tendrils no doubt may most correctly be regarded as the modified main axis which has been pushed aside by a lateral branch. The branches are then sympodia, whose successive shoots bear alternately 1 and 2 foliage-leaves: thus, on the figure there are portions altogether of 5 shoots (I.-V.), the 1-leaved ones are shaded, the 2-leaved ones are white. The following facts however are adverse to this theory: (1) the first leaf on an axillary bud is then situated 180° from the subtending leaf (e.g. the lowermost shaded leaf, Fig. 479, 180° from the lowermost white leaf), whilst the rule in the Dicotyledons is that it is placed only about 90° to one side. (2) The buds (Fig. 479 K) from which the dwarf-branches develop, must then be accessory and sister-buds to the sympodial shoots, but their first leaves have a different relative position to this, which is very peculiar, and a still more remarkable fact is that the buds, K, etc. are similar in structure and present in all the axils; thus we only find accessory buds in the cases where no tendrils are opposite to the leaves, and the main bud must then be considered to be suppressed. (3) The development proves that the tendrils arise on the side of a vigorous growing-point of the stem or by its division, and do not develop, as might be expected, from the apex of the shoot. But these relations however, find their analogues and are all capable of explanation, whereas other less natural modes of explanation are opposed to them.
435 species; especially in the Tropics; they are rarer in America. In N. Am. some Vitis-species and Ampelopsis quinquefolia are found. Vitis vinifera is supposed to have originated in the districts East and South of the Caspian Sea. Wine is obtained from Vitis-species, especially V. vinifera, and “raisins,”—(the name “currants,” given to a special variety with small, seedless fruits, is derived from Corinth).—The species of Ampelopsis (Virginian Creeper) are cultivated as ornamental plants.
Order 5. Rhamnaceæ. The stamens are placed opposite the petals as in the Ampelidaceæ (Fig. 482), but the flowers are much more perigynous or entirely epigynous. The trees and shrubs belonging to this order have simple, most frequently penninerved leaves with stipules; frequently thorny (modified branches). The flowers are inconspicuous, sometimes unisexual (Fig. 482), and have 5 (-4) sepals, petals, stamens, and generally 3 (2–5) carpels. The calyx has valvate æstivation. The petals are very small (generally less than the sepals), often spoon-like, hollow, and embracing the stamens; a disc covers the inner surface of the thalamus or the base of the style in the epigynous flower; gynœceum simple, with one style and one erect ovule in each loculus. The fruit is most frequently a drupe. The embryo is large, often green or yellow, with endosperm.
Rhamnus (Buckthorn) has a juicy drupe with 3 (2–4) stones, surrounded at the base by the persistent portion of the receptacle; the disc is thin. R. cathartica (common Buckthorn): diœcious, with opposite, serrate leaves. R. frangula (Alder Buckthorn): flowers ☿, with scattered, entire leaves.—Ceanothus (N. Am., with richly-flowered inflorescences and a fruit closely resembling that of the Euphorbias). Phylica, Pomaderris (Austr., fruit a capsule). Zizyphus, Paliurus, Colletia (S. Am.) are thorny shrubs; C. spinosa has thorny shoots with small, caducous leaves; the seedling has normal foliage-leaves. Others climb by tendrils as in the Ampelidaceæ, e.g. Gouania.
Fig. 482.—Rhamnus cathartica: A long-styled ♂-flower; pet petals; B short-styled ♂-flower; C long-styled ♀-flower; D short-styled ♀-flower (after Darwin).
475 species, 40 genera; chiefly in temperate and tropical climes. Some are medicinal plants, the bark and fruit having purgative properties (the bark of Rhamnus frangula and “Cascara Sagrada” from the bark of R. purshiana are officinal). The fruits and seeds of others are edible, for example, the fruits of Zizyphus lotus, Z. vulgaris, Z. spina Christi, etc. Green and yellow dyes are obtained from the fruit of R. cathartica, infectoria and others (Avignon grain). Ceanothus-, Rhamnus- and evergreen Phylica-species are ornamental shrubs.
Exclusively trees or shrubs with simple, entire, scattered leaves without stipules. They have a strongly perigynous, regular, 4-merous flower. The receptacle (often coloured) envelopes a simple gynœceum formed of 1 carpel and with, in most cases, 1 ovule, bearing on its edge 4 (or 5) petaloid sepals and, but rarely at the same time, small, scale-like petals. The corolla is most frequently entirely wanting (and hence these plants were formerly reckoned among the Monochlamydeæ); frequently only one of the 2 whorls of stamens, which are situated on the inner side of the edge of the receptacle, is developed. The fruit is most frequently a 1-seeded berry or drupe, or a nut which may be falsely berry-like, the partly persistent receptacle being fleshy and enveloping it.
This family appears the most nearly allied to the Frangulinæ, especially the Rhamnaceæ, and may be considered as a further development of these in the direction of the petaloid development of the receptacle and reduction of the corolla and gynœceum, which in this instance only consists of one carpel. Another deviation is that both the whorls of stamens are present, while one of these is always wanting in Frangulinæ. They also appear to be related to the Lauraceæ (see page 391).
Order 1. Thymelæaceæ. The flowers are most frequently ☿ (Fig. 483). The receptacle is high, generally tubular, coloured, and bears on its edge the 4-(or 5)-merous calyx, with imbricate æstivation. The corolla is wanting or is represented by small scales. The stamens are situated on the inside of the receptacle, and number 4 + 4 (or 5 + 5); stigma capitate. 1 pendulous ovule (Fig. 483 B), the radicle pointing upwards. The fruit is most frequently a berry. A disc is sometimes developed. Endosperm wanting or very slight.
Fig. 483.—Daphne mezereum: A flower; B longitudinal section of pistil.
Daphne (Spurge-Laurel, Fig. 483) has a deciduous receptacle, often coloured; sepals 4; petals absent; stamens 4 + 4. Berry.—Gnidia (corolla); Pimelea (2 stamens); Thymelæa; Passerina and others.
400 species; chiefly in the warm, sub-tropical zone, especially the Cape and Australia. Only Daphne and Thymelæa in Europe. In the fruit and bark of some, for example Daphne, pungent, burning and poisonous properties are found. The bark of D. mezereum (native and cultivated) and D. laureola is officinal. A specially tough bast is found in some species, for example Lagetta lintearia (Lace-tree, Jamaica), which is used in weaving. Some are cultivated in gardens as ornamental shrubs, especially species of Daphne.
Order 2. Elæagnaceæ. Shrubs or trees, which are easily recognised by the covering of peltate hairs found upon almost all parts of the plant, causing them to assume a silvery or rusty-brown appearance. Stipules are absent; the leaves are simple, most frequently scattered. Flowers (Figs. 484, 485) frequently unisexual. The sepals are valvate, 2-4; the corolla is wanting; stamens 4 + 4 or 0 + 4. The ovule is erect and the radicle turned downwards (Fig. 486). The fruit is a nut, but becomes a false fruit, being surrounded by the persistent receptacle or the lower part of it, and thus assuming a berry- or drupe-like appearance (Fig. 486). Endosperm insignificant.—Shepherdia (opposite leaves) has 4 sepals, 4+4 stamens, as in Daphne. Diœcious.—Elæagnus (Silver-leaf) is ☿, has 4–6 sepals, and 4–6 stamens alternating with them. Hippophaë is diœcious; it has 2 sepals and 4 stamens in the ♂-flower (perhaps properly speaking 2+2 stamens); thorny (stem-structures).
16 species; especially ornamental shrubs, e.g. Elæagnus argentea, angustifolia; Hippophaë rhamnoides and Shepherdia canadensis. Northern Temp.
Figs. 484–486.—Elæagnus angustifolia.
Fig. 484.—Floral diagram.
Fig. 485.—Longitudinal section through the flower.
Fig. 486.—Longitudinal section through the fruit.
Order 3 (?). Proteaceæ. This order has its chief centre in the dry regions of Australia (6/10–7/10 of about 1,000 species), a smaller number in S. Africa (2/10–3/10). a few species in S. Am. Trees or shrubs, leaves generally scattered, without stipules, and more or less dry, leathery, evergreen, and often of very different forms on the same plant (undivided, compound, etc.) The flowers are ☿ (rarely unisexual), and 4-merous in the single, petaloid perianth and in the staminal whorl; 1 carpel; sometimes zygomorphic. The perianth-leaves are generally almost free, with valvate, æstivation, often leathery. Small scales alternating with the perianth are often found at the base of the ovary. The stamens generally have extremely short filaments, and are situated opposite, sometimes quite on the tip of the perianth-leaves, in a spoon-like groove. The gynœceum is 1-locular, has 1–several ovules, and is often raised on a stalk-like internode. The fruit is a follicle or nut. The seeds, most frequently winged, have no endosperm.—Protea, Manglesia, Hakea, Banksia, Grevillea, etc. 50 genera; about 1,000 species. Several species are cultivated in our conservatories for the sake of the flowers, which are beautifully coloured and arranged in crowded inflorescences. Protandrous. It is doubtful whether they were existent in Europe in the Tertiary Period. The true systematic position of the order is doubtful. They are related to the Leguminosæ and Rosifloræ, but more closely no doubt to the two preceding orders.
The flower is generally perfect, regular and polypetalous, usually perigynous or epigynous, eucyclic and 5-merous; most frequently S5, P5, A5 + 5 or 5 + 0 and G2-5, but other numbers are found, especially 4; the flowers are very frequently obdiplostemonous. The calyx is sometimes large and the corolla small; the carpels in some are entirely free, in others more or less united. Endosperm is found in the majority. The hypogynous forms approach the Cistifloræ, the others the following families, especially the Rosifloræ. This family is not, upon the whole, so well defined and natural as most of the others. The Saxifragaceæ proper, approach very near to the Rosaceæ, especially Spiræa, and form a transition to it. The forms with opposite leaves, as Philadelphus, etc., approach the Myrtifloræ, just as the Escalloniæ appear to be closely allied to Bicornes, especially Vacciniaceæ. Finally through Pittosporaceæ, they pass over to the Frangulinæ. The family terminates in very reduced forms, on the one hand in the arborescent orders with crowded inflorescences, on the other perhaps in the very remarkable order Podostemaceæ.
Fig. 487.—Diagram of a 6-merous flower (Sedum hispanicum): w branch of scorpioid cyme in the axil of the bracteole β.
Order 1. Crassulaceæ. Nearly all are herbs or small shrubs with round, succulent branches and scattered, fleshy, often more or less round leaves, which are very rarely incised, and never have stipules. The flowers are generally borne in dichasia or unipared scorpioid cymes, which again may be arranged in racemes, umbels, etc.; they are regular, ☿, hypogynous or perigynous, and most frequently have free sepals and petals (gamopetalous corollas with sessile stamens are found in Cotyledon, Bryophyllum, Echeveria, and others); the floral formula is Sn, Pn, An + n, Gn, where n may have very different values, partly depending upon the size of the flower (e.g. 4–7 in Sedum, Fig. 487; 6–30 in Sempervivum; 4 in Rhodiola, Bryophyllum, and Kalanchoë; 5 in Echeveria, Umbilicus, Cotyledon). The carpels are free and are placed opposite the petals (Fig. 487). Fruit a syncarp composed of follicles containing many, small seeds without endosperm. Outside each carpel is found a small, nectariferous scale (Fig. 487). The northern genus, Rhodiola, is diœcious. The petal-stamens are wanting in some (Crassula, Bulliarda, and others). The floral-leaves are very often displaced upon their axillary branches. A multicarpellary gynœceum also occurs.
Sedum (Stonecrop) is generally 5-merous with 10 stamens; Sempervivum tectorum (House-leek), 12-merous, and with 24 stamens. The leaves of Bryophyllum calycinum very readily form buds, and also frequently exude water from the edges.