Fig. 367.Montia.

Diagram of flower.

Portulaca (Portulaca): flower, epigynous or semi-epigynous; fruit, a pyxidium. The stamens vary in number, and are most frequently placed in groups (in consequence of splitting) opposite the petals.—Montia: the corolla is slightly gamopetalous, but cleft on the posterior side (Fig. 367), and as a consequence of the larger size of the lateral petals, slightly zygomorphic; 3 stamens.—Calandrinia; Talinum; Anacampseros; Claytonia.

125 species; mostly in warm and temperate countries, especially the arid parts of S. Am. and the Cape. Montia fontana (Blinks) is a native plant. Portulaca oleracea is cultivated as a pot-herb in the south of Europe. A few species of Portulaca and Calandrinia are ornamental plants.

Order 7. Nyctaginiaceæ. The characteristic feature of this order is the single, regular, united, and often petaloid perianth, the lower part of which generally persists after flowering and embraces the fruit as a false pericarp. The upper portion is most frequently valvate and folded, or simply valvate in æstivation. The number of stamens varies. The free gynœceum is unicarpellate and has 1 ovule. The fruit is a nut, but becomes a false drupe, since the lower persistent portion of the perianth becomes fleshy (as in Neea, where this fleshy part is almost always crowned by the upper persistent part of the perianth. In the majority of the Mirabileæ the lower part becomes the dry anthocarp, while the upper petaloid part falls away after flowering). Finally, a peculiar involucre is formed around the flowers by free or united floral-leaves.—The majority are herbs, some are trees (Pisonia, etc.); Bougainvillea is a liane. The stems are often nodose and swollen at the nodes; the leaves are simple, penninerved, scattered, or opposite, without stipules. In some, the vascular bundles are scattered; stem anomalous.

Mirabilis; the structure of the stem is abnormal. Dichasial branching with continuation from the second bracteole, thus forming unipared scorpioid cymes. The perianth is petaloid, funnel-shaped, and has a folded and twisted æstivation resembling that of the corolla of the Convolvulaceæ; the upper coloured portion falls off after the flowering. Outside, and alternating with it, is a 5-partite, sepaloid involucre of 5 spirally-placed floral-leaves.—Oxybaphus; the involucre envelops 1–3 dichasial flowers.—Bougainvillea; the involucre is rose-coloured, 3-leaved, and envelops 3 flowers (placed laterally; the terminal flower wanting). The leaves of the involucre in Boerhaavia, Pisonia, Neea, and others are reduced to teeth or scales.

157 species; mostly in tropical countries, and especially S. Am. Species of Mirabilis (Am.) are ornamental plants. Theïn is found in Neea theïfera Oersted (discovered by Lund in Lagoa Santa, Brazil), which may be used as a tea-plant.

Order 8. Aizoaceæ. Only 3 whorls are found in the flower, which alternate with one another when their leaves are equal in number. The first is sepaloid, the third one the carpels, and the intervening one is either uncleft, in which case it is developed as stamens, or it is divided into a large number of members which then all become stamens (arranged in groups), or the outermost ones become developed as petals. The fruit is most frequently a capsule with several loculi. Most of the species are herbs with thick, fleshy stems, and exstipulate leaves. The structure of the stem is usually anomalous.

1. Aizoideæ have hypogynous or perigynous flowers with (4–) 5 perianth-leaves; stamens single, or (by splitting) in groups of 2–3, alternating with the perianth-leaves. The gynœceum (with 3–5 carpels) has 3–5 loculi in the ovary, and most frequently numerous ovules in each loculus, borne on the central placenta formed by the edges of the carpels. The fruit is a capsule. The inflorescences are dichasia and unipared scorpioid cymes.—Aizoon, Mollugo, Sesuvium, and others are herbs or bushes, most frequently hairy.

2. Mesembrianthemeæ have semi- or wholly-epigynous flowers.—Tetragonia. The perianth is 4 (more rarely 3–5–6)-merous. Stamens single, or (by splitting) in groups alternating with the perianth-leaves. There is an indefinite number of carpels, and each loculus of the ovary contains only 1 pendulous ovule. Fruit a nut or drupe. The flowers arise singly in the leaf-axils, with an accessory foliage-bud below them; in some instances there is also an accessory flower between this bud and the flower. Southern hemisphere, especially at the Cape; T. expansa, New Zealand Spinach, is a fleshy plant which is cultivated as a pot-herb (Japan, Austr., S. Am.).—Mesembrianthemum: the flowers are 5-merous; the numerous linear petals and the still more numerous stamens all arise by the splitting of 5 or 4 protuberances (primordia) alternating with the sepals. The ovary presents another characteristic peculiarity: the carpels alternating with the 5–4 stamens form an ovary (with several loculi) with the ovules at first borne, as in other cases, on the inner corner of the inwardly-turned carpels; but during the subsequent development the whole ovary is so turned round that the placentæ become parietal and the ovules assume, apparently, a position very rarely met with in the vegetable kingdom: on the dorsal suture of the carpels. Shrubs or under-shrubs, more rarely herbs with fleshy stems and simple, entire, more frequently thick or triangular leaves, containing a quantity of water. The flowers open about noon, and are brightly coloured, generally red or red-violet, but odourless. The capsules dehisce in rainy weather. 300 species, mostly found at the Cape. Some are ornamental plants. M. crystallinum (the Ice-plant) and others are covered with peculiar, bladder-like, sparkling hairs, the cell-sap of which contains salt—these serve as reservoirs of water.

Family 8. Cactifloræ.

The position of this family is very doubtful; but it seems in many respects to approach Mesembrianthemum. Some botanists place it near to the Ribesiaceæ; others, again, to the Passifloraceæ. Only 1 order.

Fig. 368.A Echinocactus: a position of a leaf-lamina; b a lateral shoot on the displaced axillary bud. B Pereskia: b a foliage-leaf on a small thorny branch which is subtended by a foliage-leaf which has fallen off and left a scar(a).

Fig. 369.Echinopsis.

Order Cactaceæ (The Cacti). The flower is epigynous, ☿, regular, and remarkable for its acyclic structure; there are, for instance, a large number of spirally-placed sepals and petals, which gradually pass over into one another, and which in some species, to a certain extent, arise from the walls of the ovary as in Nymphæa (Fig. 383 A, B). The petals are free; rotate, opening widely in Opuntia, Pereskia, and Rhipsalis; erect and united at their base into a shorter or longer tube in Cereus, Epiphyllum, Mammillaria, Echinocactus, Melocactus, and others (Fig. 369). Stamens numerous, attached to the base of the corolla; gynœceum formed of many carpels, with one style, dividing into a number of branches corresponding to the number of carpels; the ovary has one loculus with many parietal placentæ; the ovules are anatropous, on long and curved funicles. Fruit a berry with exendospermous seeds. The fruit-pulp is mainly derived from the funicles.—The external appearance of the Cactaceæ is very peculiar; Pereskia, which has thick and fleshy leaves (Fig. 368), deviates the least; foliage-leaves of the usual form are wanting in the other genera, or are usually very small, and quickly fall off and disappear (Opuntia), or are modified into thorns; the stem, without normal foliage-leaves,—so characteristic a feature in this order,—makes its appearance after the two normally developed cotyledons. The stems are fleshy, perennial, and may finally become woody. In some they are elongated, globose, pointed, and more or less dichotomously branched, e.g. in several of the Rhipsalis species, which live mostly as epiphytes on trees; in others, elongated, branched, globose, or, most frequently, more or less angular (prismatic) or grooved and provided with wings, and either columnar and erect (as much as about 20 metres in height and 1 metre in circumference, as in C. giganteus in New Mexico) or climbing by roots (Cereus and Rhipsalis-species); in others again, compressed, more or less leaf-like, often with a ridge in the centre (winged), branched and jointed: Epiphyllum, Phyllocactus, Opuntia, some species of Rhipsalis; others are thick, short, spherical or ovoid, unbranched or only slightly branched, and either studded with prominent warts (mammillæ) each of which supports a tuft of thorns (Fig. 368 A; Mammillaria and others) or with vertical ridges, separated by furrows (rows of mammillæ which have coalesced) in Melocactus, Echinocactus, Echinopsis (Fig. 369); at the same time the ovary in some is embedded in the stem so that leaves or leaf-scars, with tufts of thorns in their axils, may be observed on the ovary just as on the stem.—The flattened shoots of the Cactaceæ are formed in various ways, either by the compression of cylindrical axes (Opuntia) or, as in Melocactus, etc., from winged stems in which all the wings are suppressed except two.

The thorns are produced directly from the growing points of the axillary buds, and are modified leaves. The axillary bud is united at its base with its subtending leaf, which as a rule is extremely rudimentary; and these together form a kind of leaf-cushion, larger in some genera than in others. This leaf-cushion attains its highest development in Mammillaria, in which it is a large, conical wart (see Fig. 368 A), bearing on its apex the tuft of thorns and rudimentary lamina.—The seedlings have normal cotyledons and a fleshy hypocotyl.

All the species (1,000?) are American (one epiphytic species of Rhipsalis is indigenous in S. Africa, Mauritius and Ceylon), especially from the tropical table-lands (Mexico, etc.). Some species, especially those without thorns, as Rhipsalis, are epiphytes. Opuntia vulgaris, the fruits of which are edible, is naturalized in the Mediterranean. The cochineal insect (Coccus cacti) lives on this and some closely allied species (O. coccinellifera, etc.), particularly in Mexico and the Canary Islands. Several are ornamental plants.

Family 9. Polycarpicæ.

The flowers as a rule are ☿, regular and hypogynous; however in some orders they are unisexual, e.g. in the Myristicaceæ, or zygomorphic (in Monkshood and Larkspur in the Ranunculaceæ); in the Lauraceæ, (Fig. 386) for example, perigynous, and in Nymphæa (Fig. 383) even partially epigynous flowers are typical.—The flowers are acyclic in very many of the genera of the two first orders, if not completely so, at any rate in the numerous stamens and carpels, thus denoting an old type. It is a remarkable characteristic that in the majority of the orders the number 3 prevails in the calyx and corolla; the number 5 also occurs, but the number 2 is seldom met with. Most orders have a double perianth; chorisis does not occur, suppression is rare, and the parts of the flower are developed in acropetal succession. The most characteristic feature in the order is the free, one-leaved, as a rule numerous carpels (apocarpous gynœceum). The number of carpels in some of the last mentioned orders dwindles down to 1 (e.g. the Berberideæ and Myristicaceæ). The carpels in Nymphæaceæ become united into one pistil (syncarpous), a condition which we also find distributed among the other orders.

Endosperm occurs in almost all the orders (except e.g. Lauraceæ). The nutritive tissue in Cabombeæ and Nymphæeæ is chiefly perisperm.

Fig. 370.—Diagram of Aquilegia vulgaris: sp spur. A cyclic flower.

Fig. 371.—Diagram of a dichasium of Ranunculus acer: α1, α1, and β1, β1, bracteoles (the buds in the axils of the bracteoles, α and α1, are continued antidromously). The flower has cyclic calyx and corolla, bub acyclic (8/21) stamens.

Fig. 372.—Diagram of an acyclic Ranunculaceous flower (only 3 stamens are indicated). The spiral of the sepals has a divergence of 3/5; that of the corolla and subsequent leaves 3/8.

Order 1. Ranunculaceæ. Nearly all are herbs (except Clematis). The leaves are scattered (except Clematideæ), they have a large sheath with broad base (no stipules), and are most frequently palminerved with palmate lobes. The flowers are hypogynous, with most frequently a well pronounced convex receptacle (Figs. 374 B, 380), ☿, regular (except Delphinium and Aconitum); their structure varies very much; in some the leaves are verticillate, in others arranged spirally; in others, again, both modes of arrangement are found. It is a characteristic feature that the various series of leaves (especially calyx and corolla) are not so distinct or so sharply divided as is usual. The leaves of the perianth are free, imbricate (except Clematideæ); stamens numerous, with most frequently extrorse anthers; gynœceum free, apocarpous (except Nigella and partly Helleborus), with 1 or several ovules (Figs. 373, 378, 379) borne on the ventral suture. The fruit is either a nut or a follicle (Actæa has berries). The seed has a large, oil-containing endosperm and a small embryo (Fig. 374).

The main axis generally terminates in a flower, and the lateral axes branch in a cymose manner (Fig. 371). The flowers show the following differences in construction: VERTICILLATE (EUCYCLIC), i.e. constructed all through of alternating whorls: Aquilegia (Fig. 370), Xanthorhiza, and sometimes Eranthis. Semiverticillate (HEMICYCLIC) i.e. with sepals and petals in alternate whorls, and the others arranged spirally: Ranunculus (Fig. 371), Myosurus, Pæonia and several other genera entirely, or in certain species only. Spiral-flowered (ACYCLIC) i.e. all the leaves are arranged spirally, so that sepals and petals do not alternate the one with the other, even though they are the same in number: Adonis (Fig. 372), Aconitum, Delphinium-species, Nigella-species, Helleborus. The leaves of the calyx are in this instance arranged on a spiral of 2/5; those of the corolla on 2/5, 3/8, 5/13 or 8/21, and stamens and carpels likewise on higher fractions of the same series.

The genera Caltha, Anemone, Thalictrum and Clematis have a single perianth, which is most frequently petaloid; it is thus apparent that the sepals are petaloid, and the leaves, which in other genera have developed as petals, are in these instances stamens. The calyx is similarly petaloid in the genera Helleborus, Eranthis, Nigella, Delphinium and Aconitum; but the petals are present in these instances in unusual (horn-like) forms, and almost entirely given up to the function of nectaries, a function they already possess in Ranunculus. According to a more recent theory the “honey-leaves” are transformed stamens, which have lost the function of reproduction; the perianth is then single, and most frequently petaloid. [Those leaves in the flowers of many Ranunculaceæ which bear nectaries are termed by Prantl honey-leaves, and comprise those leaf-structures of the flower whose essential function lies in the production of nectar, and which, independent of the differentiation of the perianth into calyx and corolla, are derived from the stamens by the loss of their reproductive functions. Clear transitional forms are found between the two series of the perianth (e.g. between the sepaloid and petaloid perianth-leaves of Anemone japonica, A. decapetala, Trollius-species) while transitional forms are never found between perianth-and honey-leaves (with the exception of Aquilegia vulgaris, var. stellata). In Anemone and Clematis the honey-leaves pass gradually into the stamens, and agree with the stamens in the other Ranunculaceæ in their arrangement, development, and scant system of veins (except Nigella). In Delphinium, sect. Consolida, the two honey-leaves placed in front of the unpaired perianth-leaf are united into one, as shown by the veins (twice three veins arranged symmetrically). The honey-leaves of Aquilegia, Callianthemum, and the majority of the Ranunculus-species serve by reason of their large circumference, as organs of attraction, and on this account are considered as petals by other authors.—The same position in the flower which the honey-leaves assume is found occupied by staminodes, without nectar, in some Coptis-species, in Anemonopsis, Actæa sect. Euactæa, (e.g. A. racemosa), Clematis sect. Atragene; in the last-named they closely surround the stamens, in Actæa they are petaloid.—A perianth, sharply differentiated into calyx and corolla, and destitute of honey-leaves, is found in Anemone, sect. Knowltonia (Cape),

Adonis, Pæonia.—The perianth of the Ranunculaceæ is considered by Prantl to be usually petaloid.—The nectaries arise in the Ranunculaceæ (1) on normal stamens (Clematis sect. Viorna), (2) on the honey-leaves (this is generally the case), and (3) on the carpels (Caltha and the majority of Trollius-species).—As the result of his researches upon the Ranunculaceæ, Prantl does not agree with the view advanced by Drude (Schenk, Hand. d. Bot. iii.) that the petals in general have proceeded from the metamorphosis of the stamens (K)].

Fig. 373.—Ovaries in longitudinal section: v the ventral suture; d the dorsal suture: A, B Clematis; C Ranunculus; D Myosurus.

The most primitive form of fruit is undoubtedly the pod formed by one carpel, on the edges of which (along the ventral suture) two rows of ovules are situated: Pæonieæ, Helleboreæ, Delphinieæ (Fig. 379). In a great many genera the number of ovules has been limited to one perfect one, which is placed in the central plane under the united leaf-edges, and sometimes also some barren ovules above it (Fig. 373). The fruitlets in this case become achenes, and are present in much larger numbers than when there are follicles.

Fig. 374.Helleborus niger: A flower; B receptacle; pet petals (honey-leaves); pi stamens and carpels; C seed; D anther (cross section); alb endosperm.

Fig. 375.Caltha palustris: fruit.

The following have Follicles: Pæonieæ, Helleboreæ (except Actæa) and Delphinieæ; Achenes: Ranunculeæ, Anemoneæ and Clematideæ.

A. Follicles (Figs. 375, 379), with many ovules, situated in two rows along the ventral suture. Actæa has berries, Nigella has capsules of several loculi.

1. Pæonieæ, Peony Group. This has regular, acyclic flowers with a normal, most frequently 5-leaved, imbricate calyx; large, coloured petals, and introrse anthers. Slightly perigynous. Surrounding the base of the carpels a ring-like swelling of the receptacle (“disc”) is present, which is largest in P. moutan. The follicles are more or less fleshy or leathery. Mostly herbs, with pinnatisect or decompound leaves and large, solitary flowers; a gradual transition may be traced from the foliage-leaves to the petals. Pæonia; Hydrastis.

Fig. 376.Aquilegia vulgaris.

Fig. 377.Caltha palustris (nat. size).

Fig. 378.Nigella: A, B fruit of N. damascena, entire, and cut transversely. C Petal (honey-leaf) of N. arvensis. D Petal of N. damascena.

2. Helleboreæ, Hellebore Group. This has regular flowers with most frequently a coloured calyx. The petals (honey-leaves) are modified into nectaries; they may be horn-like, provided with a spur, or of a similarly unusual form, or they may be entirely absent. Anthers often extrorse.—Trollius (Globe-flower[36]). The flower is acyclic: many petaloid sepals, succeeding these, most frequently, several linear, dark yellow petals, which bear a naked nectary at the base; finally, many stamens and carpels arranged in a spiral (3/8, 8/21).—Caltha (Marsh-marigold, Figs. 375, 377); 5 (-7) yellow sepals, no petals. The foliage-leaves have a large amplexicaul sheath.—Helleborous (Hellebore) has pedate leaves. The flower is acyclic, with 5 large, regular, persistent, often petaloid sepals (2/5); small, horn-like petals (honey-leaves; most frequently 13, divergence 8/13) and generally few carpels (Fig. 374).—Coptis.Isopyrum.Eranthis (Winter Aconite), like Anemone, has a 3-leaved involucre and most frequently trimerous flowers, 6 large petaloid sepals, 6 petals (tubular honey-leaves), 6 oblique rows of stamens, 3–6 carpels. Aquilegia (Columbine, Fig. 376); the flower is entirely cyclic and has large spurs on all the 5 petals (funnel-shaped honey-leaves); S5 coloured, P5, A5 × (8–12), G5 in regular alternation (Figs. 376, 370); the innermost stamens are often staminodes (Fig. 370).—Nigella (Love-in-the-mist, Fig. 378) has 5 sepals and 8 small, two-lipped petals cleft at the apex (the nectary is covered by the under-lip; Fig. 378 C, D). The 5 carpels are more or less completely united; and a many-carpellate ovary with free styles is formed in some. Large air-chambers in the external wall of the ovary are formed in N. damascena (Fig. 378).—Actæa (Baneberry) has coloured sepals, either no petals or an indefinite number, and only 1 carpel. The fruit is a berry (or follicle).—Cimicifuga, Garidella, Xanthorhiza (S5, P5, A5 + 5, G5).

3. Delphinieæ, Larkspur Group. Zygomorphic flowers with coloured calyx; the 2 posterior petals (honey-leaves) are transformed into nectaries, the others are small or absent altogether.—Aconitum (Monkshood); 5 sepals, of which the posterior one (Fig. 379 A) is helmet-shaped; most frequently 8 petals (as in Fig. 372), of which the two posterior ones (honey-leaves) are developed into long-clawed nectaries (Fig. 379 A, k) enveloped by the helmet-like sepal; the others are small, or are to some extent suppressed. Stamens on a spiral of 3/8–5/13; generally 3 carpels. Perennial herbs.—Delphinium (Larkspur); very closely allied to Aconitum, but the anterior 4 petals are most frequently wanting, and the 2 posterior ones have each a spur, which is enclosed by the posterior sepal, the latter being also provided with a membranous spur. Stamens and carpels arranged on a spiral of 3/8, 5/13, 8/21. In D. ajacis and consolida there is apparently only 1 petal (by the fusion of 4) and 1 carpel.

Fig. 379.Aconitum napellus. A Flower in longitudinal section, below are the 2 bracteoles; a half of helmet-like sepal; b and c other sepals; k nectary; f carpels. B Ovary in longitudinal section; C the same transversely; d dorsal suture; v ventral suture.

B. Fruit achenes. Many carpels, each with only 1 ascending (Fig. 373 C), or pendulous (Fig. 373 D), perfect ovule; often also rudimentary ovules above it (Fig. 373 A, B). Fruit achenes.

4. Ranunculeæ, Buttercup Group, has double perianth. Myosurus and Adonis have pendulous ovules as in Anemoneæ (Fig. 373 D); Ranunculus, with Batrachium and Ficaria, erect ovules (Fig. 373 C) and downwardly-turned radicle.Ranunculus. Most frequently S5, P5, many spirally-placed stamens and carpels (Figs. 371, 380). The petals (honey-leaves) have a nectary at the base, covered by a small scale. Batrachium, Water Ranunculus, deviates by the achenes being transversely wrinkled; dimorphic leaves. Ficaria has 3 sepals and 7–8 petals arranged in 2/5–3/8. F. ranunculoides (the only species) has tuberous roots, which spring from the base of the axillary buds, and together with these, serve as organs of reproduction. The embryo has only 1 cotyledon.Myosurus (Mouse-tail) has small prolongations from the 5 sepals; 5 narrow petals which bear the nectaries near the apex; sometimes only 5 stamens, and an ultimately very long receptacle, with numerous spirally-arranged achenes (Fig. 381).—Adonis is acyclic (Fig. 372); most frequently 5 sepals with a divergence of 2/5, 8 petals of 3/8, indefinite stamens and carpels of 3/8 or 5/13. The corolla has no nectary.

Fig. 380.—Flower of Ranunculus sceleratus in longitudinal section.

Fig. 381.Myosurus minimus: c cotyledons; m the foliage-leaves; f the floral axis with the carpels, and g the same without; y insertion of perianth.]

5. Anemoneæ, Anemone Group, has a single perianth. (Pendulous ovules (Fig. 373 D), radicle turned upward).Anemone has a single, petaloid, most frequently 5–6-leaved perianth, and beneath the flower most frequently an involucre of 3 leaves, placed close together in the form of a whorl. In A. nemorosa, ranunculoides, etc., the involucral leaves resemble foliage-leaves; in A. hepatica they are situated close under the perianth, and resemble sepals, and in the sub-genus Pulsatilla they stand between the foliage-leaves and floral-leaves. The style of Pulsatilla finally grows out in the form of a feather. The main axis of A. hepatica has unlimited growth (it is biaxial), and the flowers are borne laterally in the axils of the scale-leaves; in the others (uniaxial) the flower is terminal, and the rhizome becomes a sympodium after the first flowering.Thalictrum (Meadow Rue) has no involucre; 4–5-leaved, greenish perianth. The receptacle is flat. The stamens are brightly-coloured and have long filaments; 1–5 accessory flowers may occur in the leaf-axils of the panicle-like inflorescence.

6. Clematideæ, Clematis Group. This differs from all the others in the valvate æstivation of the calyx and its opposite leaves. There are 4 (-several) petaloid sepals; petals are absent, or linear (Atragene). Ovule 1, pendulous. Achenes, often with prolonged, feathery style. The majority of the genera are shrubs, and climb by their sensitive, twining leaf-stalks.—Clematis; Atragene.

Pollination. The flowers are conspicuous either by coloured petals (honey-leaves) (Ranunculus, Pæonia) or coloured sepals (Helleborus, Anemone, Caltha, etc.), or by both (Aquilegia, Delphinium), or by the coloured stamens (Thalictrum). Some have no honey (Clematis, Anemone, Thalictrum), and are generally visited by insects for the sake of their pollen. Others have nectaries on the corolla (Ranunculus, Trollius, Helleborus, Nigella, Aconitum, etc.), more rarely on the stamens (Pulsatilla, Clematis-species), or the carpels (Caltha), or the calyx (certain species of Pæonia). The honey is readily accessible in the flat, open flowers, and these flowers also may easily pollinate themselves. There is marked protandry where the honey lies deeply hidden, as in Aquilegia, Delphinium, and Aconitum. Helleborus and some Ranunculus-species are protogynous.

About 680 species; especially in northern temperate climates, and extending to the Polar and Alpine regions. Only the Clematideæ are tropical.

The order has an abundance of acrid, vesicant properties (R. acer, sceleratus, etc.), and poisonous alkaloids (Helleborus niger is poisonous). Officinal: Aconitum napellus (aconitine; leaves and tuberous roots); the rhizome of Hydrastis canadensis from N. Am. (the alkaloid hydrastine). The order, however, is best known for its ornamental plants; almost all the genera have species which are cultivated for their beauty. Sweet-scented flowers are absent.

Order 2. Nymphæaceæ (Water Lilies). Water Plants; generally with large, floating leaves, and large solitary flowers; sepals 3–5, petals 3–∞, stamens 6–∞, carpels 3–∞. The flower is hypogynous, but in the Nymphæeæ different degrees of epigyny are found, and from this fact, as well as from the carpels being united into one pistil, the family forms a lateral offshoot from the Ranunculaceæ, with much greater modification. The seed often has an aril, and, in the majority, a farinaceous nutritive tissue, partly endosperm, partly perisperm (Fig. 383 C). The embryo has 2 thick cotyledons and a small hypocotyl; the plumule is well developed, with 2–4 leaves.

1. Cabombeæ. 3–4 species (Tropical S. Am.), resembling the Water Ranunculus, with two kinds of leaves, the submerged being dissected and the aerial peltate. The flowers are eucyclic, trimerous, with 2–3 free, epigynous carpels. The ovules are situated on the central line of the carpel—an almost unique circumstance. Endosperm and perisperm. Cabomba; Brasenia.

2. Nelumboneæ. The leaves are peltate, raised on long stalks high above the water. Large, hypogynous flowers (Fig. 382); sepals 4–5; petals numerous; stamens numerous; carpels several, distinct. The receptacle is very remarkable, being raised above the stamens, and developed into an inverted conical body on the apex of which the nut-like fruits are embedded in pits. Endosperm is wanting, but the embryo is large and has well developed cotyledons.—Nelumbo, 2 species. N. lutea (N. Am.); N. speciosa (E. Ind.) was sacred amongst the ancient Hindoos and Egyptians, (the Lotus flower); its seeds are used as food.

3. Nymphæeæ, Water Lily Group. The carpels are united into one, many-locular ovary, whose numerous ovules are situated on the surface of the partition walls (as in the Poppies); the stigma is sessile and radiating, the number of rays corresponding to the number of carpels (Fig. 383). The fruit is a spongy berry with many seeds, which have a large perisperm in addition to the endosperm (Fig. 383 C).

Fig. 382.Nelumbo nucifera: vertical section through the receptacle.

Sepals, petals, and stamens often pass gradually over the one into the other, the petals becoming narrower by degrees, and bearing anthers on each side of the apex, which gradually become larger anthers in proportion to the filament, until the perfect stamen is developed. The long-stalked leaves are floating, and most frequently cordate, elliptical, leathery, with a shiny surface, sometimes (as in Victoria regia and Euryale ferox) with strongly projecting thorny ribs on the lower surface. In the intercellular passages of the leaves are some peculiar, stellate cells.

Nuphar has 5 sepals, and an hypogynous flower. The petals, which are small, have a nectary on the back; the coloured inner side of the sepals functions as petals; the ovate gynœceum is quite free.—N. luteum is a native plant (Yellow Water-Lily), with, most frequently, 13 petals and 10–16 loculi in the ovary. The rhizome is horizontal, as much as 5–6 cm. in thickness, and bears on its under surface a number of roots, which on dying-off leave deep scars; the leaves are borne in spiral lines, and the flowers are solitary in certain leaf-axils. The construction of the rhizome is very peculiar; the vascular bundles are scattered and closed as in a monocotyledonous stem.

Fig. 383.Nymphæa: A flower in longitudinal section, the most external leaves being removed; B fruit; C seed of Nuphar (longitudinal section); the perisperm at the base, the endosperm at the top surrounding the embryo.

Nymphæa has 4 sepals, and the flower is more or less epigynous. Petals and stamens are inserted at different heights on the ovary to just beneath the stigma (Fig. 383). Nymphæa alba (White Water-Lily). Victoria regia from the Amazon, and Euryale ferox from Asia, have entirely epigynous flowers. The shield-like leaves of Victoria are as much as 2 metres in diameter, and the edge is bent up to a height of 5–14 cm.; the flowers are 20–40 cm. in diameter, and change in twenty-four hours from white to rose-red. A development of heat, as much as 14°C. above the temperature of the air, together with a strong formation of carbonic acid, has been observed during flowering.

Pollination. Nymphæa alba and other species of the sub-genus Symphytopleura are self-pollinated; the sub-genus Leptopleura is insect-pollinated. Nuphar and Victoria can effect self-fertilisation; Euryale is self-fertilised, often in entirely closed and submerged flowers.—The dissemination of the seeds in Nuphar luteum is effected by the fruit, which rests on the water, becoming detached from its stalk, and dehiscing from the base upwards so that the seeds are set free; while in Nymphæa alba the spirally-twisted stalk draws the fruit under water, and it dehisces by its upper part being thrown off as a hood, and the seeds which are enclosed in air-tight sacs rise to the surface of the water. In this condition they are able to float and can only sink to the bottom when the air has disappeared.

53 species; in fresh water in all parts of the world, but especially in the Tropics.—The rhizomes and seeds of some may be used as food; Euryale ferox is even cultivated. Nymphæa cœrulea and Lotus were sacred among the Egyptians.

Order 3. Ceratophyllaceæ. About 3 species. Aquatic plants, submerged, rootless; leaves cartilaginous, verticillate, dissected into repeatedly dichotomous branches which are finely toothed; only one of the leaves in a whorl supports a vegetative branch. The flowers are monœcious, axillary. Inside the 6–12 perianth-leaves are situated in the ♂-flower 10–20 stamens with thick connective, and in the ♀-flower a gynœceum formed by one carpel, with one orthotropous and pendulous ovule, which has only one integument. Fruit a nut, which, in some species, bears on each side a pointed horn, and at the apex a similar one, formed by the persistent style.—The embryo has an unusually well developed plumule with several whorls of leaves. The plant is rootless throughout its whole life.—Ceratophyllum (Horn-wort).

Order 4. Annonaceæ. Sepals 3; petals 3+3 (most frequently valvate); succeeding these (as in the Ranunculaceæ) are numerous acyclic stamens and an apocarpous gynœceum; the flowers are hypogynous, regular and ☿, generally very large (2–3 cm. in diameter), and the leaves of the perianth are more or less fleshy or leathery. The majority have syncarps with berry-like fruitlets, but in Annona and some others the carpels fuse together into a large, head-like fruit—a kind of composite berry. The seeds have ruminate endosperm as in Myristica.—Trees or shrubs with alternate, simple, entire, penninerved leaves without stipules. 450 (700?) species; especially tropical. The best known are Anona cherimolia, squamosa and reticulata (all from America) cultivated on account of their large, delicious fruits. Some have acrid and aromatic properties (Xylopia, Cananga—the flowers of the latter yield Ylang-ylang); Artabotrys odoratissimus; Asimina (N. Am.).

Order 5. Magnoliaceæ. Trees or shrubs with scattered, often leathery, entire leaves, generally with stipules, which (as in Ficus) are rolled together and form a hood round the younger internodes above them, and are cast off by the unfolding of the next leaf, leaving a ring-like scar. The endosperm is not ruminate. Corolla imbricate. Fruit a syncarp.

A. Magnolieæ. The flowers are borne singly, and before opening are enveloped in an ochrea-like spathe which corresponds to the stipules of the foliage-leaves. The perianth generally consists of 3 trimerous whorls, the external one of which is sometimes sepaloid (Liriodendron, and the majority of Magnoliaspecies), sometimes coloured like the others; the perianth is sometimes many-seriate. Numerous spirally-placed stamens and carpels. The latter are situated on the elongated, cylindrical receptacle, and are individually more or less united, except in Liriodendron, where they are free. This last genus has winged achenes; the fruitlets in Magnolia open along the dorsal and ventral sutures, and the seeds then hang out, suspended by elastic threads formed from the vascular bundles of the funicle and raphe; they are red and drupaceous, the external layer of the shell being fleshy—a very rare occurrence.

B. Illicieæ has no stipules. The carpels are situated in a whorl on a short receptacle. Follicles, one-seeded. The leaves are dotted by glands containing essential oil. Illicium; Drimys.

70 species; in tropical or temperate climates; none in Europe or Africa. They are chiefly used as ornamental plants, e.g. the Tulip-tree (Liriodendron tulipifera, N. Am.), Magnolia grandiflora (N. Am.), M. yulan and fuscata (China), and others. The remains of Liriodendron occur as fossils in the Cretaceous and Tertiary periods.—The fruits of Illicium anisatum (Star-aniseed from Eastern Asia) are OFFICINAL. The bark of Drimys winteri (S. Am.) is also strongly aromatic.

Order 6. Calycanthaceæ. These are very closely related to the Magnoliaceæ, but differ in having perigynous flowers with many perianth-leaves, stamens and (about 20) carpels in a continuous spiral, seeds almost devoid of endosperm with rolled up, leaf-like cotyledons, and leaves opposite on a square stem.—There are some species in N. America (Calycanthus florida, occidentalis, etc.) and 1 in Japan (Chimonanthus præcox), all strongly aromatic.

Order 7. Monimiaceæ. Aromatic shrubs with opposite leaves. Perigynous flowers. The anthers dehisce by valves like those of the Lauraceæ, and the Monimiaceæ may thus be considered as an apocarpous form of this order. They are also closely related to Calycanthaceæ. 150 species, tropical.—Hedycarya, Mollinedia, Monimia.