A handbook of systematic botany

Part 35

Chapter 353,020 wordsPublic domain

=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._

=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.~

=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.

=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_).

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.~

_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 _Magnolia_species), 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._

Order 8. =Berberidaceæ (Barberries).=--The regular, ☿, hypogynous flowers are dimerous or trimerous and have regularly alternating whorls of free sepals, petals, and stamens and 1 unilocular carpel; the corolla and stamens have each 2 whorls, the calyx at least 2. The anthers open, as in Lauraceæ, by (2) _valves_, but are always introrse (Fig. 384). The pistil has a large, disc-like, almost _sessile_ stigma (Fig. 385), and in the ovary _several_ erect ovules are placed close to the base of the ventral suture. The fruit is most frequently a _berry_. Seeds endospermous.--Shrubs or herbs with scattered, most frequently compound leaves (without stipules), and racemose inflorescences.--~They show a relationship to the Lauraceæ in the number of the parts of the flower and the dehiscence of the anthers.~

_Berberis_ is a shrub; it has sepals 3 + 3, petals 3 + 3, stamens 3 + 3 (Fig. 384). The petals (honey-leaves) bear internally at the base 2 darkish-yellow nectaries. The filaments are sensitive at the base, and suddenly bend inwards if touched at that spot (Fig. 385). ~The racemes often have a terminal, 5-merous flower; they are borne on dwarf-branches. The leaves on the long-branches develope into thorns, but the buds in their axils, in the same year as themselves, develope as the short-branches with simple foliage-leaves, _articulated_ at the base, from which fact some authorities have considered that the leaf is compound with a single, terminal leaflet.~--_Mahonia_ has imparipinnate leaves. The flower has 3 whorls of sepals. Otherwise as in _Berberis_.--~_Epimedium_; herbs with spurred petals; the flowers dimerous; 4–5 whorls of sepals, 2 of petals and stamens. Fruit a capsule. _Leontice_, fruit dry. The anthers of _Podophyllum_ dehisce longitudinally.--_Nandina. Aceranthus._~

100 species; North temp., especially Asia: fossils in Tertiary. _Berberis vulgaris_ is a native of Europe. This and other species, together with _Mahonia aquifolium_ (N. Am.), _Epimedium alpinum_, etc., are cultivated as ornamental plants. Several have a yellow colouring matter in the root and stem. OFFICINAL: the rhizome of _Podophyllum peltatum_ (from N. Am.) yields podophyllin.

Order 9. =Menispermaceæ.= This order has derived its name from the more or less crescent-like fruits and seeds. Diœcious. The flowers are 2–3-merous, most frequently as in _Berberis_ (S3 + 3, P3 + 3, A3 + 3), with the difference that there are 3 _free carpels_, each with 1 ovule; in some genera, however, the number is different. Stamens often united into a bundle (as in _Myristica_); anthers dehiscing longitudinally; fruit a drupe.--The plants (with herbaceous or woody stems) belonging to this order are nearly all _twining_ or _climbing_ plants, and have scattered, palmate or peltate, sometimes lobed leaves without stipules. Structure of stem anomalous. _Cocculus, Menispermum, Cissampelos, Anamirta._

150 species; Tropical; very rich in bitter and poisonous properties. OFFICINAL: Calumba-root from _Jateorhiza columba_ (E. Africa). The following are cultivated as ornamental plants:--_Menispermum canadense_ (N. Am.) and _M. dahuricum_ (Asia). The fruits of _Anamirta cocculus_ (E. Ind.) are very poisonous (“Grains-of-Paradise”; the poisonous matter is picrotoxine).

Order 10. =Lardizabalaceæ.= This order, by the free, apocarpous carpels, belongs to a more primitive type, and by the united stamens to a more developed one. _Akebia_; _Holbœllia_; principally climbing or twining shrubs. About 7 species in S.E. Asia and S. Am.