A handbook of systematic botany
Part 28
The flowers in this family are _hypogynous_ and have in part the general monocotyledonous type with 5 trimerous whorls completely developed in a regular hermaphrodite flower, and in part the flowers so much reduced that the type is very difficult to trace. On the one hand the family is well developed and has capitate inflorescences (_Eriocaulaceæ_) and on the other hand it is distinctly reduced (_Centrolepidaceceæ_). This family has taken its name from the fact that the ovule is not, as in the Liliifloræ and nearly all other Monocotyledons, anatropous, but _orthotropous_, so that the embryo (βλάστη) becomes placed _at the end of the seed opposite_ (ἐναντίος) _to the hilum_. Large, mealy endosperm.--The orders belonging to this family are by certain authors grouped with the _Bromeliaceæ_ and _Pontederiaceæ_, etc., into one family, FARINOSEÆ, so named on account of the mealy endosperm, the distinguishing character of the Liliifloræ then being that the endosperm is fleshy and horny.
Order 1. =Commelinaceæ.= The complete Liliaceous structure without great reductions in the number of whorls, but with generally few ovules in each loculus of the ovary, is found in the Commelinaceæ, an almost exclusively tropical order with about 317 species; herbs, some of which are introduced into our gardens and greenhouses. The stems are nodose; the leaves often _clasping_; the flowers are arranged in unipared scorpioid cymes, often so that they form a zig-zag series falling in the median line of the bracts, and after flowering they bend regularly to the right or left, outwards or inwards. They are more or less _zygomorphic_, particularly in the stamens, which in the same flower are of different forms or partially suppressed. The outer series of the _perianth_ is sepaloid, the inner petaloid, generally violet or blue; the filaments are sometimes clothed with hairs formed of rows of bead-like cells (well known for showing protoplasmic movements). Fruit a trilocular _capsule_ with loculicidal dehiscence (generally few-seeded); in some a nut. The radicle is covered by an external, warty, projecting covering which is cast off on germination.--The abundant raphides lie in elongated cells whose transverse walls they perforate.--_Commelina, Tradescantia, Tinantia, Cyanotis, Dichorisandra_.
Order 2. =Mayacaceæ.= This order is closely allied to the Commelinaceæ. 7 species. American marsh- or water-plants.
In many of the following orders of this family the flowers are united into compound inflorescences, with which is accompanied a reduction in the flower.
Order 3. =Xyridaceæ= (50 species). Marsh-plants with radical, often equitant leaves arranged in 2 rows, and short spikes on long (twisted) stalks. The flowers, as in the Commelinaceæ, have sepals (which however are more chaffy) and petals, but the outer series of stamens is wanting. Capsule (generally many-seeded).
Order 4. =Rapateaceæ.= Marsh-plants with radical leaves, usually in two rows, and several spikelets on the summit of the main axis, clustered into a capitulum or unilateral spike. Each spikelet has numerous imbricate floral-leaves and one flower. 24 species. South America.
Order 5. =Eriocaulaceæ.= The “Compositæ among Monocotyledons,” a tropical order. The flowers are borne in a _capitulum_ surrounded by an _involucre_, very similar to that of the Compositæ. The flowers are very small, unisexual, ♂ and ♀ often mixed indiscriminately in the same capitulum; they have the usual pentacyclic structure; the leaves of the inner perianth are often connate and more membranous than the outer; in some the outer series of stamens are suppressed; in each of the 3 loculi is one pendulous ovule. Capsule. The leaves are generally radical and grass-like.--335 species; _Eriocaulon_, _Paepalanthus_, etc., _E. septangulare_ on the west coast of Scotland, and Ireland, and in North America.
Order 6. =Restiaceæ.= A small, especially S. African and S. Australian, xerophilous order (about 235 species), which is quite similar in habit to the Juncaceæ and Cyperaceæ. The leaves are often reduced to sheaths. The flowers are diœcious, the perianth as in _Juncus_, but the outer series of stamens suppressed. The ovary and fruit as in Eriocaulaceæ; the ovary, however, may be unilocular, and the fruit a nut. _Restio_, etc.
Order 7. =Centrolepidaceæ.= These are the most reduced plants in the family; small grass- or rush-like herbs. The flowers are very small, naked. Stamens 1–2, carpels 1–∞. 32 species. Australia.--_Centrolepis_ (flowers generally ☿ with 1 stamen and 2–∞ carpels).
Family 5. =Liliifloræ.=
The flower is constructed on the general monocotyledonous type, with 5 alternating, 3-merous whorls (Fig. 278), but exceptions are found as in the Iridaceæ (Fig. 279) by the suppression of the _inner_ whorl of stamens; in a few the position in relation to the bract differs from that represented in Fig. 278, and in some instead of the trimerous, di- or tetramerous flowers are found (_e.g. Majanthemum_, _Paris_). Flowers generally _regular, hermaphrodite_, with simple, _petaloid_, coloured perianth (except, for example, Bromeliaceæ); ovary trilocular, generally with 2 ovules or 2 rows of ovules in the inner angle of each loculus (Fig. 304 _C_, _D_). _Endosperm_ always present.--A very natural family, of which some divisions in part overlap each other. The habit varies; the leaves are however long, entire, with parallel venation, except in Dioscoreaceæ (Fig. 313).
In the first orders of this family the flowers are hypogynous, and in the first of all the styles are free, and the capsule dehisces septicidally; in the following the flowers are epigynous and in some reduced in number or unisexual; capsule with loculicidal dehiscence, or a berry.
HYPOGYNOUS flowers: Colchicaceæ, Liliaceæ, Convallariaceæ, Bromeliaceæ (in part).
EPIGYNOUS flowers: Amaryllidaceæ, Iridaceæ, Bromeliaceæ (in part), Dioscoreaceæ.
Order 1. =Colchicaceæ.= The flower (Fig. 304 _A_) is ☿, regular, _hypogynous_, trimerous in all five whorls (6 _stamens_); anthers usually _extrorse_. Gynœceum with 3 _free styles_ (_A, D_); fruit a _capsule with septicidal dehiscence_ (_E_); embryo very small (_F_). The underground stem is generally a corm or rhizome, seldom a bulb.
=A.= VERATREÆ.--_Veratrum_; perennial herbs, stem tall with long internodes and broad, folded leaves; the flowers andromonœcious, with free, widely opening perianth-leaves (Fig. 304 _A_), and globular anthers; inflorescence a panicle.--_Zygadenus, Melanthium, Schœnocaulon, Uvularia, Tricyrtis_.
=B.= TOFIELDIEÆ.--_Narthecium_ and _Tofieldia_ have leaves alternate (arranged in two rows), sword-like and borne in rosettes; racemes or spikes. _Narthecium_ forms an exception to the order by having a simple style and fruit with loculicidal dehiscence; _Tofieldia_ by the introrse anthers. In this they are related to the Liliaceæ. _Narthecium_ has poisonous properties, like many other Colchicaceæ.
=C.= COLCHICEÆ.--_Colchicum_ (Autumn Crocus); perennial herbs, with a long, _funnel-shaped, gamophyllous perianth_, and introrse anthers. The flowers of _C. autumnale_ spring up immediately from the underground stem, which is in reality a _corm_ formed of one internode. ~_Colchicum autumnale_ flowers in autumn without leaves; in spring the radical foliage-leaves appear simultaneously with the fruit. The flower is protogynous, and is pollinated by insects (humble-bees, etc.) which seek the honey secreted by the free part of the stamen a little way down the tube. The length of the tube protects the fruit, and not, as in other cases, the nectary.--_Bulbocodium_ and _Merendera_ have unguiculate perianth-leaves, free, but closing together like a tube.~
175 species; chiefly in North America and South Africa. _Tofieldia_ is an Arctic plant. The order is rich in pungent, poisonous alkaloids (veratrin, colchicin, etc.). OFFICINAL; the seeds of _Colchicum autumnale_ (Europe) and _Schœnocaulon officinale_ (Mexico), and the rhizome of _Veratrum album_ (mountains of Central Europe).
Order 2. =Liliaceæ (Lilies).= Flowers as in the Colchicaceæ but with _introrse_ anthers; _ovary free, 3-locular, with single style; capsule_ 3-locular with _loculicidal_ dehiscence.--The majority are herbs with _bulbs_; the inflorescence is _terminal_. In many species reproduction takes place by means of bulbils (small bulbs) formed in the axils of the foliage-leaves (_e.g. Lilium bulbiferum_, _lancifolium_, etc., _Gagea lancifolia_, etc.), or in the bracts of the inflorescence (many species of _Allium_); in many species several buds are developed as bulbs in the axils of the bulb-scales themselves (accessory buds arising close together), and in some the formation of buds is common on the leaves.
=A.= TULIPEÆ, TULIP GROUP. Bulbs. The aerial, elongated stem bears the foliage-leaves. Flowers few but generally large, with free perianth-leaves. _Tulipa_; style absent, no honey; flowers generally solitary, erect.--_Fritillaria_ perianth campanulate with a round or oblong nectary at the base of each perianth-leaf.--_Lilium_; perianth widely open, generally turned back with a covered nectary-groove in the centre of each segment. Anthers versatile.--_Lloydia; Erythronium._
=B.= HYACINTHEÆ, HYACINTH GROUP. Bulbs. Leaves radical; aerial stem leafless with raceme or spike. In some the perianth-segments are free, in others united. Honey is produced often in glands or in the septa of the ovary (septal glands).--_Ornithogalum_ has a leafy stem; _Scilla_; _Eucomis_ has a tuft of floral-leaves above the raceme; _Agraphis_; _Hyacinthus_; _Puschkinia_; _Chionodoxa_; _Muscari_; _Veltheimia_; _Urginea_.
=C.= ALLIEÆ, ONION GROUP. Generally bulbs. Leaves radical. Stem leafless with a compound umbellate or capitate inflorescence of unipared helicoid cymes, which before flowering are surrounded by two broad involucral leaves.--~_Allium._ Filaments often petaloid and bidentate; in many species bulbils are found in the inflorescence.--Some species have flat leaves: _A. sativum_, Garlic; _A. porrum_, Leek; _A. ursinum_; others have round, hollow leaves: _A. cepa_, Onion; _A. fistulosum_, Winter Onion; _A. ascalonicum_, Eschalot; _A. schænoprasum_, Chive.~--_Gagea_; honey is secreted at the base of the perianth, no special nectary; inflorescence few-flowered.--_Agapanthus; Triteleia._
=D.= ANTHERICEÆ. Rhizome; raceme; the leaves not fleshy and thick.--_Anthericum_; _Asphodelus_; _Bulbine_; _Chlorophytum_; _Bowiea_ has an almost leafless stem with curved, climbing branches.
=E.= ALOINEÆ, ALOES. Stem generally aerial and tree-like, bearing on its summit thick, fleshy leaves, often with a thorny edge (Fig. 306). Raceme branched or unbranched.--_Aloë_; _Gasteria_; _Yucca_ (has secondary thickening, p. 274).
=F.= HEMEROCALLIDEÆ. _Phormium_, (_Ph. tenax._ New Zealand Flax); _Funckia_ (_Hosta_); _Hemerocallis_.
At this point the following are best placed: _Aphyllanthes_ (_A. monspeliensis_); _Xanthorrhæa_ (Black-boy); _Xerotes_; _Lomandra_; _Kingia_; the very membranous, dry perianth of the last resembles that of the Juncaceæ, and also there are only 1–few ovules in the loculi.
POLLINATION by insects. Honey in some is produced on the perianth (see Tulipeæ), in others by glands on the carpels (in the septa and parietal placentæ, septal glands): _Hyacinthus_, _Allium_, _Anthericum_, _Asphodelus_, _Yucca_, _Funckia_, _Hemerocallis_, etc. Some _Allium_-species are protandrous. _Fritillaria_ is visited by bees, _Lilium martagon_ by moths, _L. bulbiferum_ by butterflies, _Phormium_ (New Zealand) by honey-birds.
About 1,580 species; rare in cold climates; their home is in sunny plains with firm, hard soil, and warm or mild climate, particularly in the Old World (S. Africa; As. Steppes; Mediterranean); at the commencement of spring the flowers appear in great profusion, and after the course of a few weeks disappear; during the hot season their life lies dormant in the bulb, hidden underground. The woody species are tropical.--The majority of the _introduced_ Liliaceæ (_Fritillaria imperialis_, Crown-imperial; _Lilium candidum_; _Tulipa gesneriana_; Hyacinth; _Muscari_-species; _Scilla_-species; _Ornithogalum nutans_; _Hemerocallis fulva_ and _flava_; _Asphodelus luteus_ and _albus_) come from the Mediterranean and W. Asia; _Funckia_ from China and Japan; several Lilies from Japan and the Himalayas; _Agapanthus_ from the Cape; _Allium sativum_ is a native of the Kerghis-Steppes; _A. cepa_ from Persia (?); _A. ascalonicum_ is not known wild (according to others a native of Asia Minor), perhaps a form of _A. cepa_; _A. schænoprasum_ from the N. temp. region.
Many bulbs have pungent properties; many Onions are used as culinary plants. The bast fibres of _Phormium tenax_ (New Zealand Flax) are used technically. Dyes are obtained from the _Aloe_; gum for varnish from the stem of _Xanthorrhæa hostile_ and _australe_. OFFICINAL; “Aloes,” the dried sap of S. African species of _Aloe_ (_A. Africana_, _A. ferox_, etc.); the bulb known as “Squills” from _Urginea_ (_Scilla_) _maritima_ (Mediterranean).
Order 3. =Convallariaceæ.= This order differs from the Liliaceæ in having the _fruit a berry_ (Fig. 308) and _in never being bulbous_; the seeds are less numerous.
=A.= CONVALLARIEÆ, LILY OF THE VALLEY GROUP. Rhizome (Fig. 307) and normal foliage-leaves.--_Polygonatum_: rhizome creeping; aerial shoot leafy, bearing the flowers in racemes in the axils of the foliage-leaves; perianth tubular. _P. multiflorum_ (Solomon’s seal), _P. officinale_, etc.--_Majanthemum_: flower 2-merous; perianth almost polyphyllous, spreading. _Smilacina. Streptopus_ (_S. amplexifolius_; the flowers or inflorescence unite with the entire succeeding internode).--_Convallaria_ (1 species _C. majalis_, Lily of the valley); flowers in terminal racemes; 2 basal foliage-leaves; perianth globose, bell-shaped. _Reineckea carnea_ (Japan, China) in gardens.--_Paris_ (_P. quadrifolia_, Herb-Paris); flowers solitary, terminal, 4-merous, polyphyllous; styles 4, free (approaching the Colchicaceæ; it is also poisonous); a whorl of 4 (-more) 3-nerved, reticulate leaves on each shoot.--Ornamental plants: species of _Trillium_, _Aspidistra elatior_ (Japan).
=B.= ASPARAGEÆ, ASPARAGUS GROUP. Scale-like leaves and green assimilating branches.--_Asparagus_: horizontal rhizome. The aerial shoots are very richly branched; the numerous needle-like bodies upon the plant are _leafless shoots_, which are crowded together in double scorpioid cymes in the axils of the scale-leaves; the two first lateral axes, placed outside to the left and right, generally bear flowers. Polygamous.--~_Ruscus_ (Butcher’s broom) is a S. European _shrub_ with _leaf-like_, ovoid or elliptical shoots (phylloclades) which are borne in the axils of scale-like leaves, and bear flowers on the central line. Diœcious. Stamens 3, united, anthers extrorse. _Semele androgyna_ bears its flowers on the edge of the flat shoot.~
=C.= SMILACEÆ. _Smilax_ (Sarsaparilla) (Fig. 308); _climbing_ shrubs with the leaf-sheath produced into tendrils. The leaves have 3–5 strong nerves proceeding from the base, and are reticulate. Orthotropous or semi-anatropous ovules. Diœcious (Fig. 308 _C_, _E_).
=D.= DRACÆNEÆ. Fruit in some a berry, in others a capsule. The stem of DRACÆNA, when old, has the appearance of being dichotomously branched; it has the power of increase in thickness, and may become enormously thick. The Dragon-tree of Teneriffe, measured by Humboldt, attained a circumference of 14 m. and a height of 22 m.; the leaves are large, linear or linear-lanceolate.--_Cordyline_ (East Asia), various species in gardens and greenhouses (_Yucca_ is closely allied). _Astelia._
POLLINATION. _Paris quadrifolia_ and _Convallaria majalis_ have no honey, and are chiefly visited by pollen-collecting bees (in the absence of insect visits self-pollination takes place); _Polygonatum multiflorum_ has honey secreted by septal glands and protected by the base of the tubular perianth; it is pollinated by humble-bees, etc. _Asparagus officinalis_ has small, polygamous, greenish, honey-bearing flowers; the ♂-flower is almost twice as large as the ♀; both have rudiments of the opposite sex.
About 555 species; especially from N. America, Europe, and Central Asia.
OFFICINAL: “Dragons’-blood,” a red resinous juice from the stem of _Dracæna_ and the roots of some Central American species of _Smilax_. The tuberous stems of the Eastern Asiatic _Smilax glabra_ are officinal. The flowers of _Convallaria majalis_ have been lately used as a substitute for _Digitalis_. Pungent, poisonous properties are possessed by _Paris_. None of the species are used as food, except the young annual shoots of _Asparagus officinalis_, a shore-plant which is used as a vegetable.
Order 4. =Pontederiaceæ.= Flowers generally zygomorphic, hypogynous, ☿, with handsome, white or violet, petaloid perianth which forms a tube at its base. The stamens are inserted at different heights in the perianth-tube, and are reduced to three (in _Heteranthera_ seldom to one). In some the ovary is trilocular with ∞ ovules (_Eichhornia_), in others reduced to one loculus with one ovule (_Pontederia_). Fruit a capsule or nut. Embryo as long as the abundant, mealy endosperm.--Tropical water-plants (22 species) with peculiar sympodial branching, nearly the same as in _Zostera_. Spikes without floral-leaves. Many intercellular spaces in the stem and leaf.--In greenhouses: _Eichhornia azurea_, _E. crassipes_ (both from tropical and sub-tropical S. America); the latter has swollen petioles which serve as floats and enable it to float freely on the water, sending down its roots into the mud. _Heteranthera reniformis, H. zosterifolia. Pontederia cordata._
Order 5. =Amaryllidaceæ (Narcissi).= The flower is _epigynous_, otherwise exactly the same as in the Liliaceæ (6 stamens). The majority, like these, are also _perennial_ herbs with bulbs and scapes. The fruit and the other characters as in the Liliaceæ. The external appearance is, however, very different.
=A.= AMARYLLEÆ have bulbs and the leaves generally arranged in two rows; the flowers are borne singly or in umbel-like inflorescences on lateral scapes, while the main axis of the bulb is unlimited. Beneath the inflorescence is an _involucre_ (Fig. 309).--_Galanthus_, Snowdrop, has a polyphyllous perianth without corona; the three inner perianth-leaves are emarginate and shorter than the outer; the anthers dehisce apically. ~_Leucojum_ differs in having the perianth-leaves equal in length.--_Amaryllis_ has a funnel-shaped perianth, entirely or nearly polyphyllous, but somewhat zygomorphic. _Crinum; Hæmanthus; Clivia._~--_Narcissus_ has a tubular _corona_, a ligular structure arising from the perianth-tube exterior to the outer stamens. ~In _Pancratium_ (Fig. 309) the corona is united with the filaments which appear to spring from its edge. _Eucharis amazonica._~
=B.= HYPOXIDEÆ. The leaves, which are grass-like, dry, folded, and in some hairy, spring from a rhizome, generally with a divergence of 1/3. Flowers small, perianth polyphyllous, persistent, on which account perhaps the Hypoxideæ may be considered as the least altered type. The chief characteristic is that the embryo is separated from the hilum. _Hypoxis_; _Curculigo_ (_C. recurvata_, a favourite ornamental plant; S.E. Asia).
=C.= ALSTRŒMERIEÆ. (_Alstrœmeria_, _Bomarea_); stems long, leafy, often climbing.
=D.= VELLOSIEÆ (_Vellosia_, _Barbacenia_); stem woody, usually dichotomously branched, with terminal, single flowers; it bears numerous aerial roots which pierce the leaves and surround the stem. Stamens often (by splitting) 6–18. High table-lands of S. America and S. Africa.
=E.= AGAVEÆ. Very similar to the Bromeliaceæ both in their distribution (nearly all American) and in external appearance. They appear as gigantic bulbous plants with perennial, aerial, generally short stem, and perennial, large, lanceolate or linear, stiff, thick, and often thorny leaves, which form a large rosette; after the course of several (8–20) years the terminal inflorescence is developed, which is 10–12 m. high, paniculate, and freely branched. Before the inflorescence expands, a large quantity of sugar-containing sap is collected from _A. americana_ by removing the terminal bud; this on distillation yields “pulque,” the national drink of Mexico. After flowering the entire shoot dies, but the subterranean lateral shoots survive and reproduce the plant.--_Agave americana_, etc.; _Fourcroya_; _Polianthes tuberosa_ (Tuberose; Central America).
DISTRIBUTION. The 650 species are chiefly natives of S. Africa and S. America. _Clivia_, _Hæmanthus_, _Amaryllis_ are from the Cape; _Narcissus_ from S. Europe, whence many species have been introduced; _Galanthus_ and _Leucojum_ are especially from S. and Central Europe, and from the Caucasus.
USES, few, except as ornamental plants: _Galanthus nivalis_; _Leucojum_; _Narcissus pseudonarcissus_, _N. poeticus_, _N. jonquilla_, _N. tazetta_, etc.; _Amaryllis_, _Alstrœmeria_, _Eucharis_, _Crinum_, _Vallota_, etc. The vascular bundles of the various species of _Agave_ (_Agave rigida_, var. _sisalana_, sisal hemp,) are used for cordage, etc.
Order 6. =Bromeliaceæ.= The flowers are hypogynous, epigynous or semi-epigynous; the perianth is divided into _calyx_ and _corolla_; stamens 6. The fruit is a capsule or berry with many seeds. Endosperm _mealy_, embryo small, at the edge of the endosperm, but not enclosed by it.