Alliaceae

K. Rahn

Alliaceae J. Agardh, Theoria Syst. PI.: 32 (1858), nom. cons. Liliaceae subfam. Allioideae Engl. (1887). Amaryllidaceae tribus Allieae Hutchinson (1934). Milulaceae Traub, Plant Life 28:129 (1972).

Acaulescent or short-stemmed biennial or perennial geophytes; alliaceous odour often present; stem usually swollen and often forming a bulb, tuberous rhizome or rarely a corm, enveloped by sheathing, dry leaves or leaf bases. Leaves linear, filiform, lanceolate or rarely ovate, flat, angular, terete, or fistular, often fleshy, forming closed sheaths below, and sometimes forming a pseudostem around the scape; veins parallel. Scape from apex of each shoot or bulb, sometimes also lateral, terete, flat or angular, sometimes fistular, leafless except at apex. Inflorescence usually umbel-like, formed of 1 or more contracted helicoid cymes, rarely reduced to a single flower, in one species a spike (Milula). Inflorescence subtended by 2 or less frequently 1 or several, membranous, sometimes ± united spathe bracts, enveloping the young inflorescence, its branches or individual pedicels sometimes also subtended by. smaller membranous bracts. Pedicels not articulate. Flowers hermaphrodite, usually actino-morphic, zygomorphic in Solaria, Miersia, and Gilliesia. Tepals 3+3 (rarely 3 or 3+2), usually similar and petaloid, united at base, almost free, or forming a tube; tepal lobes erect, spreading or recurved. Scales or appendices between tepals and stamens, of different origin, often forming a corona. Functional stamens usually 3+3, rarely 3 or 2 and then the missing ones often transformed to staminodes. Filaments inserted on the tepals, free from each other or united, often with lateral, dorsal or apical appendices. Anthers versatile, basifixed, introrse, opening with longitudinal slits. Ovary superior or in Allium siculum and A. tripedale almost semiinferior; tricarpellary, trilocular; provided with septal nectaries; 2-several ovules in each locule. Ovules anatropous or campylotropous, bitegmic. Style solitary, erect, at apex of ovary or in Allium and Milula ± gynobasic. Stigma capitate or trilobate with a Dry or sometimes Wet (Leucocoryne) surface. Fruit a loculicidal capsule with few-numerous seeds. Seeds either rather small, ovoid or ellipsolidal to subglobose (rounded in transection), or larger, angular, semiovoid to semiglobose (triangular in transection), or in Tulbaghia flat. Testa with a crust of phytomelan. Endosperm with fatty oils and aleuron, but no starch. Embryo short and straight, or long and curved. Vegetative buds produced instead of flowers in some species of Allium.

Comprising 13 genera and about 600 species; most genera in S America, particularly Chile, the genus Allium with 260-700 species in the northern hemisphere.

Vegetative Morphology. The leafy part of the stem is short in all species. Each shoot ends in an inflorescence, and shoot structure is sympodial. Most species (except Tulbaghia and some Allium) have a true bulb (Fig. 27); the stem is short, often depressed conical and covered by one or several swollen sheathing leaves or leaf bases, externally with dry leaf bases. The new main bulb appears from the axil of the uppermost leaf; increase bulbs originate in the axils of lower leaves in the old bulb. The scape is often apparently lateral, because the green leaves spring from a lateral shoot or bulb that is to flower the next year. In Tulbaghia the stem or rhizome is more or less swollen and irregularly shaped, tuberous, covered by dry, fibrous leaves; the closed sheath formed at the base is very short. In the rhizome-bearing species of Allium the rhizome is covered by thin and fibrous leaf bases only and may be the only storage organ. The leaf bases often form long, closed sheaths around the younger leaves and the scape (Fig. 27, 28). The increase bulb in Allium (Mann 1960) produces a single contractile root which pulls the increase bulb deeper into the soil away from the mother bulb (Fig. 12F). Leaves are distichous, at least in Milula and some species of Allium (e.g. A. ampeloprasum), or else apparently spirally arranged.

Vegetative Anatomy. Lacticifers have been found in all Alliaceae screened by Sterling and Huang (1972) (Allium, Nothoscordum, Tristagma and Tulbaghia). Raphides of calcium oxalate are recorded as being absent in Allium, Milula and Tulbaghia. The stomata are anomocytic. Epicu-ticular waxes are arranged in non-orientated platelets (several Allium) or in threadlike, reticulate projections (Ipheion uniflorum and Nothoscordum bivalve); neither the Convallaria type nor the Strelitzia type was found (Frölich and

Milula Allium
  1. 27A-H. Alliaceae. A, B Allium altaicum. A Habit. B Tepals and stamens. C, D A. polyrhizum. C Habit. D Tepals and stamens, the stamens of inner whorl conspicuously appendaged. E, F A. setifolium. E Habit. F Tepals and stamens. G A. funkiifolium, habit. H A. oleraceum, inflorescence with flowers and bulbils. (Takhtajan 1982)
  2. 27A-H. Alliaceae. A, B Allium altaicum. A Habit. B Tepals and stamens. C, D A. polyrhizum. C Habit. D Tepals and stamens, the stamens of inner whorl conspicuously appendaged. E, F A. setifolium. E Habit. F Tepals and stamens. G A. funkiifolium, habit. H A. oleraceum, inflorescence with flowers and bulbils. (Takhtajan 1982)

Barthlott 1988). The veins in cylindric leaves, and often also in flat leaves in Allium, are arranged in a ring with the phloem closest to the epidermis. The cylindric leaf is probably primitive in Allium (unlike, e.g. in Juncus).

Inflorescence Structure. The scape is apical in the shoot (bulb). In some species lateral inflorescences also appear. The inflorescence in

Anatomy Dyed Shooty Leaf

Fig. 28A-I. Alliaceae. A, B Allium schubertii . A Habit. B Opened perianth with stamens. C, D A. verticillatum. C Habit. D Fruit. E-G A. paradoxum• E Habit. F Dehiscing capsule. G Seed with elaiosome. H, I A. chamaemoly. H Habit. I Capsule. (Takhtajan 1982)

Transmission Tissue Pistil
  1. 29A-G. Alliaceae. Pistil organisation in Alliaceae. A-C Transversal sections at the level of the compitum. D-G Longitudinal sections through a plane of symmetry. Pollen transmission tissue dotted; septal nectaries cross-hatched. Note differences in style insertion, opening of nectaries, and shape of placentae. A, D Nothoscordum borbonicum. E Allium nigrum. B, F A. asperum. C, G A. ampeloprasum. (Di Fulvio 1973)
  2. 28A-I. Alliaceae. A, B Allium schubertii . A Habit. B Opened perianth with stamens. C, D A. verticillatum. C Habit. D Fruit. E-G A. paradoxum• E Habit. F Dehiscing capsule. G Seed with elaiosome. H, I A. chamaemoly. H Habit. I Capsule. (Takhtajan 1982)

Nothoscordum, Allium and probably also in the other genera is formed of one or more contracted helicoid cymes, which are spikelike in Milula and umbel-like in all other genera, but sometimes reduced to 1 or 2 flowers. The inflorescence is subtended by usually 2, less frequently 1 (Milula, some Allium) or more (Nothoscordum fictile and

  1. 29A-G. Alliaceae. Pistil organisation in Alliaceae. A-C Transversal sections at the level of the compitum. D-G Longitudinal sections through a plane of symmetry. Pollen transmission tissue dotted; septal nectaries cross-hatched. Note differences in style insertion, opening of nectaries, and shape of placentae. A, D Nothoscordum borbonicum. E Allium nigrum. B, F A. asperum. C, G A. ampeloprasum. (Di Fulvio 1973)
  2. sessile) membranous bracts, which envelop the young inflorescence. The spathe bracts sometimes sheath at base, and the 2 sheaths are sometimes more or less united. The short branches in the inflorescence or the individual pedicels are sometimes also subtended by smaller membranous bracts (most species of Tulbaghia, many Allium and a few Nothoscordum). In the inflorescence of some species of Allium the flowers may partly be replaced by bulbils.

Flower Structure. Septal nectaries are probably present in all species (recorded in nine Allium and two Nothoscordum species by Daumann 1970). According to Di Fulvio (1973), the opening of the nectary is close to the style in Nothoscordum and close to the base of the ovary in Allium. A very short gynophore may be found in some Nothoscordum species (Fig. 29).

Acording to Daumann (1970), a cuticula is present in the septal nectaries of Nothoscordum, but not in those of Allium. A nectarium parenchyma is present, and vascularisation absent in the species studied.

Embryology. In Nothoscordum, embryo-sac development follows the Polygonum type, while in Allium and Leucocoryne the megaspore mother cell undergoes only the first meiotic division and the chalazal dyad cell develops into an bisporic Allium-type embryo sac (Davis 1966).

In the facultatively parthenogenetic Allium ramosum the synergids and antipodal nuclei are supposed to fuse in pairs and to give rise to supernumerary embryos: apogamy. Adventitious em-bryony from nucellar tissues is recorded several times from Allium, Nothoscordum borbonicum and probably other related species (Schnarf 1931; Guaglianone 1972). In contrast to Agapanthus, a parietal cell is not cut off from the archesporial cell in Allium, Leucocoryne, Miersia, Nothoscordum and Tulbaghia (Wunderlich 1959). The endosperm is Nuclear in Allium and Helobial in Nothoscordum and Tulbaghia. The anther tape-tum is secretory; the tapetum cells uninuclear in 4 species of Allium, and 2-4-nuclear in Tulbaghia as in most other genera in Asparagales (Wunderlich 1954). Microsporogenesis is successive, and the pollen 2-celled when dispersed.

Pollen Morphology. Schulze (1980), who studied the pollen of eight genera (Tulbaghia, Allium, Nothoscordum, Ipheioti, Tristagma, Solaria, Miersia and Gilliesia) found it invariably sulcate and reticulate. The mean length of the pollen is between 30 and 45 |xm, and the sulcus does not reach the proximal side of the pollen grain.

  1. Most species of Tulbaghia have 2n = 12 (Vosa 1975). Milula and most species of Allium have 2n = 16 (but also 2n = 32,14,28 and 18 are found in Allium). In Nothoscordum (Nunez et al. 1974; Crosa 1975; Ravenna 1978) five species have 2 n = 10, two 2 n = 16, two 2 n = 18, two 2 n = 19, and one 2n = 26. In this genus acrocentric chromsomes are about 1/2 the length of the metacentric chromosomes. The sum of the number of acrocentric and twice the number of metacentric chromosomes are multiples of 8. Alternatively, the chromosomes can be considered combinations of sets with 4 metacentric and sets with 3 metacentric + 2 acrocentric (Crosa 1981). Four species are diploid, six tetraploid and one hexaploid when a set is considered "the basic number". Two Ipheion species were studied by Crosa (1975); they had chromosomes similar to, but smaller than Nothoscordum, 2n = 20 in I. sessile (4 metacentric and 16 acrocentric) and 2n = 12 in J. uniflorum (2 metacentric and 10 acrocentric). Tristagma, the other genus with which Nothoscordum has been associated, was studied by Crosa (1981). Three species have 2n = 8 (6 meta-, 2 acrocentric); one 2n = 16 (12 meta- and 4 acrocentric); one unidentified species 2n = 24 (18 meta- and 6 acrocentric). The chromosomes are of a size similar to those of Nothoscordum. In Leucocoryne narcissoides Ravenna (1978) counted 2n = 24, and 2n = 12 or 18 has been recorded in other species of the same genus. In Miersia chilensis 2n = 12,20 and 21 were recorded. The chromosome numbers in the other genera are unknown.
  2. All or most species are pollinated by insects. Beetles, flies, bees and butterflies are recorded as pollinators of Allium. Flowers of most species of Allium are protandrous, a few are protogynous. Insects are attracted by the usually showy mass of flowers and by the nectar, which is produced in septal nectaries, and in Allium siculum and A. tripedale in 3 glands at the base of the inner filaments. Some species of Allium (e.g. A. rotundum, A. sphaerocephalum) have cleisto-gamous flowers and autogamy.

Fruit and Seed. The fruit is a loculicidal capsule, with 2-several seeds in each of the 3 locules. Seeds are flat and elongate with a straight embryo in Tulbaghia. Seeds in the other genera are angular and more or less isodiametric, 1.7-4 mm long. Allium often has seeds with a long embryo, which is curved at 120-180°, or the embryo is shorter and then curved at less than 90°. In the other genera the seeds have a short and thick embryo. An eleiosome may be present (Allium triquetrum). The epidermis of the testa has a crust of phytomelan 10-28 jim thick in Allium and Milula, thinner in the other genera. The epidermis cells are isodiametic or slightly elongate. The inner layers of the testa are compressed or collapsed, as is the tegmen. Endosperm without starch, but with aleuron and fat.

Dispersal. Seeds of some Allium species are dispersed by ants (e.g. A. ursinum and A. triquetrum).

Reproductive Biology. Vegetative propagation is largely restricted to the formation of increase bulbs. In some species of Allium, such as A. vineale, small inflorescence propagules (bulbils) are produced at the base of the pedicels or from transformed gynoecia. Apospory has been recorded for Allium and in Nothoscordum borbonicum and is probably also present in other related species (Guaglianone 1972).

Phytochemistry (Hegnauer 1963, 1986). Characteristic for Alliaceae is the absence of alkaloids, which are regularly found in Amaryllidaceae. Saponins are recorded from Allium, Nothoscordum, Leucocoryne, Tristagma and Gilliesia, and not recorded as being absent in any species of Alliaceae. The scales of onions are particularly rich in flavonoids (quercetin glycosides) and therefore are used for dyeing Easter eggs, textiles, etc. Starch has not been found in bulbs of Allium and is replaced by several other carbohydrates, including galactane, raffinose and the inulinelike fructane. Starch is found in all bulbs or rhizomes studied from the other genera (Nothoscordum, Ipheion, Tulbaghia). Calcium oxalate is not present as raphides in Allium, Milula or Tulbaghia, but as solitary crystals or other conglomerations in Allium (Jaccard and Frey 1928) and Nothoscordum (Guaglianone 1972: 193). The characteristic alliaceous odour is present in Milula-, in some or most species of Tulbaghia, Allium, Leucocoryne, Ipheion and Tristagma; in one Gilliesia; but noted neither in any species of Nothoscordum, Schickendantziella, Speea, Trichlora, Solaria, nor Miersia. This odour is released from wounded or decaying tissue only, when alliin, an S-substituted cysteine derivative, under the influence of the enzyme alliinase, is degraded to the water-soluble and strong-smelling allicin, pyruvic acid and ammonia. Alliin has four homologues, allylalliin, propylalliin, methylalliin and ethylalliin, of which methylalliin is widely distributed among plants (e.g. in all Brassicaceae and within various other families). Ipheion uniflorum contains methyl-, propyl-, and ethylalliin. The lachrymatory factor from Allium cepa is propanthial-S-oxide derived from propenylsulhuric acid and is liberated from onion cells on crushing; it is almost absent in Allium ampeloprasum "Porrum" and A. sativum. All these sulphur compounds are potent feeding deterrents to animals.

Subdivision and Relationships Within Alliaceae. Krause (1930) divided his Liliaceae subfam. Alloideae into four tribes: Agapantheae with Agapanthus and Tulbaghia; Miluleae with Milula; Gilliesieae with Trichlora, Speea, Erinna, Solaria, Miersia, Gethyum, Gilliesia and Ancruma, i.e. our genera nos. 9 and 11-14. The tribe Allieae contained Allium, Nothoscordum, Tristagma, Steinmannia, Leucocoryne, and all genera now in Themidaceae.

Traub (1963) divided his Amaryllidaceae subfam. Allioideae into four tribes: Miluleae (with Milula), Allieae, Gilliesieae (with Schickendantziella, Speea, Miersia, Gethyum, Gilliesia, Solaria, Erinna, Trichlora and Ancruma, i.e. our genera nos. 7, 8 p.p. and 9-13); and Agapantheae (with Agapanthus). Allieae were divided into three subtribes: Alliinae (with Allium, Nectaroscordum, Caloscordum, Steinmannia, Nothoscordum, Leucocoryne, Tulbaghia, Stemmatium and Tristagma, i.e. our genera nos. 1-6 and 8 p.p.), Brodiaeinae (here in Themidaceae) and Millinae (here in Themidaceae).

After a parsimony analysis of rbcL sequences, Fay and Chase (1996) proposed placing Agapanthus in Amaryllidaceae subfam. Agapanthoideae. Alliaceae were divided into three subfamilies: Allioideae (with Allium and Milula); Tul-baghioideae (with Tulbaghia only); and Gilliesioideae with the rest of the genera here placed in Alliaceae.

Affinities. After the rejection of Gagea (Schnarf 1948), the Alliaceae (including Themidaceae and Agapanthus) were long considered an indisputable monophyletic taxon. The superior ovary placed it in Liliaceae, until Hutchinson (1934), mainly because of the umbellate inflorescence, placed it in Amaryllidaceae. Dahlgren et al. (1985) placed Alliaceae in Asparagales together with Hyacinthaceae, Amaryllidaceae and 28 other families. Alliaceae were separated from Amaryllidaceae by having steroidal saponins, superior ovary (like most other Asparagales), and absence of alkaloids. Based on a parsimony analysis of rbcL sequences, Fay and Chase (1996) found that "Alliaceae sensu Dahlgren et al. were poly-phyletic, with Agapanthus sister to Amaryllidaceae, and Brodiaeeae (with the exception of Petronymphe) being more closely related to Hyacinthaceae ..., than to the Amaryllidaceae-Alliaceae complex".

Distribution and Habitats. The majority of the Alliaceae species are found between 25° and 45° latitude N and S, Tulbaghia in southern Africa, Milula in Himalayas, Allium in N America, Europe, N Africa and especially the drier parts of Asia. The rest of the genera are found in S America, especially in Chile and Argentina.

The Alliaceae probably evolved in adaptation to a long dry season; but a few species are (probably secondarily) adapted to a more humid climate, e.g. Allium ursinum to woods in Europe.

Parasites. Cultivated Allium are attacked by many parasites. The larva of leek moth (Acrolepia assectella) eats especially the young leaves of Allium ampeloprasum "Porrum". The larva of the onion fly (Delia antiqua) destroys the bulbs or the young plants of Allium. The onion thrips (Thrips tabaci) does much harm, mainly by transferring virus. Leaves and scapes of several species are attacked by rust (Puccinia porri), and the young plants by a smut (Urocystis cepulae). Peronospora destructor attacks the leaves and scapes, and Botrytis allii the bulbs, especially of A. cepa.

Economic Importance. The cultivated species of Allium have been treated by Moore (1954/55). Allium cepa "Cepa", onion, is the most commonly cultivated and used as a vegetable or for flavouring food. A. cepa "Aggregatum", shallot (for which the name A. ascalonicum has been misapplied), is used in a similar way or pickled. Less common is A. fistulosum, Welsh onion, and A. chinen$e> rakkyo. The leaves of A. schoenoprasum, chive, and A. tuberosum, Chinese chive, are used for flavouring food, as are the increase bulbs (cloves) of A. sativumy garlic. A. ampeloprasum, "Porrum", leek, has a milder taste and is used as a vegetable. Some species of Allium are used as ornamentals. Less frequently, Tulbaghiay Notho-scordum> Ipheion and Leucocoryne are used for the same purpose.

Key to the Genera

1. The closed sheaths at base of leaves very short; distinct bulb structures absent; rhizome or tuber containing starch, covered by dead leaves; inflorescence umbellate; flowers actinomorphic; stamens 6, included in the perianth tube; corona fleshy, annular or of 3-6 scales in the throat; seeds many, flat; embryo straight or slightly curved, phytomelan crust rather thin. Alliaceous odour present. S Africa 1. Tulbaghia

- The closed sheath at base of leaves usually long; rhizome or bulb with or without starch; flowers actinomorphic or zygomorphic; stamens 2-6, free or included in the perianth tube; a corona present or absent; seeds few or many, more or less isodiametric and angular, phytomelan crust thin or relatively thick. Alliaceous odour present or absent. Northern hemisphere and S America 2

2. Style ± gynobasic, young ovary with the style in a depression at the summit; bulb without starch; flowers actinomorphic; tepals 6; 6 anthers fertile; ovules few, rarely several per locule; most species with alliaceous odour; embryo often long and curved. Northern hemisphere 3

- Style not in a depression at top of young ovary, bulbs with starch; few species with alliaceous odour; usually several ovules in each locule; tepals 3-6, 2-6 stamens fertile; embryo straight, usually short. S America 4

3. Inflorescence umbellate or flowers 1 or 2; alliaceous odour present or absent. N Africa, Europe, Asia, N America

2. Allium

- Inflorescence spicate of many flowers, when young covered by 1 large spathe; with alliaceous odour. Himalayas, Tibet 3. Milula

4. Six stamens fertile 5

- Less than 6 stamens fertile 11

5. No corona or scales between stamens and tepals; tepals united below 6

- Staminodes, scales or corona between perianth and fertile stamens 10

6. Spathe bracts free from each other or united for less than 25%; stamens free from each other or united below, in 1 or 2 series; flowers solitary or several; testa smooth to the naked eye 7

- Spathe bracts 2, united with each other at 1 margin for more than 75%, the other for more than 30%. Lower 3050% of tepals united. Stamens free from each other, inserted in the tube in 2 series; flowers usually solitary; testa pitted. S Brazil, Uruguay, Argentina, Chile 5. Ipheion

7. Tepals united below for 10-65%; filaments free or slightly united below 8

- Tepals almost free; filaments united, forming a tube, enveloping the ovary 9

  1. Lower 10-30% of tepals united, anthers protruding, filaments in one series, sometimes united with each other below. S America, one species cosmopolitan
  2. Nothoscordum

- Lower 25-65% of tepals united, anthers usually included into the tepal tube; filaments free from each other, in 2 series. Chile, S Argentina 6. Tristagma

  1. Tepals 3; stigma entire. NW Argentina
  2. Schickendantziella

- Tepals 6; stigma slightly 3-lobed. Chile 8. Speea

10. Flowers actinomorphic; corona in throat; anthers in tepal tube 9. Leucocoryne

- Flower zygomorphic; 6 scales between staminal tube and linear tepals. Chile 12. Miersia

11. Three outer stamens fertile, 3 staminodes or coronar appendices; filaments free or absent 12

- Three or 2 anterior stamens fertile; the posterior are small, staminodial, or absent; filaments united 13

12. Inner and outer tepals almost equal; staminodes long, protruding from the throat; some species with alliaceous odour. Chile 9. Leucocoryne

- Inner tepals short, obtuse, scalelike; outer lanceolate, almost free; staminodes small. Peru 10. Trichlora

13. Corona absent or of 6 small violet scales; filaments forming a tube below. S Chile, S Argentina 11. Solaria

- Corona of irregular scales on the outside of the staminal cup; staminal tube split down one side. Chile

  1. Gittiesia
  2. Tulbaghia L.

Tulbaghia L., Mant. 148, 223 ("Tulbagia") (1771), orth. et nom. cons.; Vosa, Ann. Bot. (Rome) 34:47-121 (1975), rev.; Burbidge, Notes R. Bot. Gard. Edinb. 31:77-104 (1978), rev.

With alliaceous smell. Rhizome, corm or bulb with thick roots. Old dried or slightly fleshy leaf bases enveloping the rhizome or corm. Leaves 4-8. Flowers 3-40. Spathe bracts 2, enclosing the floral buds. Bracts subtending the individual pedicels. Perianth 7-14mm long, forming a tube for about half its length, corona of 3-6 scales or forming a fleshy tube. Anthers in 2 whorls, sessile in the tube formed by the perianth or the corona. Style short. Stigma capitate. Seeds several in each cell. The phytomelan crust thin. Embryo in the mature seed almost straight, x = 6, most species diploid. Twenty two spp., S Africa.

Allium L., Sp. PL: 294 (1753); Steam, Ann. Mus. Goulandris 4:

83-198 (1978), Europ. spp.; Pastor 8( Valdés, Rev. genero

Allium Peninsula Ibérica y Islas Baleares (1983); de Wilde-

Duyfjes, Belmontia 7: 1-237 (1977), African spp. Nectaroscordum Lindl. (1836). Caloscordum Herbert (1844).

A large number of other synonyms have been described but never generally recognised.

Most species with alliaceous odour and a tuni-cated bulb. Leaves basal, but their sheathing bases may form a flat or terete, sometimes fistular pseudostem. Spathe bracts 1-2, in some species also individual bracts at base of each pedicel. Inflorescence umbel-like with few to many flowers, sometimes with bulbils. Tepals 6, small, free or slightly united below, very rarely with more than 1 nerve. Stamens 6, all or rarely only 3 fertile, usually attached to the base of the tepals. Filaments often dilated at the base. Septal nectaries open in lower part of ovary, or openings absent. Ovules 1-10 in each locule, often 2. Seeds broad and triangular in transection, with a thick phytomelan crust. 2 n = 16 in most species. Estimated 260-550-690 spp., N Africa, Europe, Asia and N America, especially abundant in Central Asia (Flora USSR has 225 species).

Subdivisions controversial. Allium siculum Ucria and A. tripedale Trautv. from SE Europe and W Asia with semiinferior ovary, discoid receptacle or the pedicel expanded above, and 2n = 18 have been referred to the genus Nectaroscordum. Allium neriniflorum (Herb.) Baker and A. tubiflorum Rendle from E Asia without alliaceous odour, tepals united below and forming a cup, have been placed in a separate genus Caloscordum or in Nothoscordum. Several species economically important, see above under the family.

3. Milula Prain

Milula Prain, Ann. R. BoL Gard. (Calcutta) 5: 164 (1896).

With alliaceous odour. Leaves linear. One large monophyllous spathe bract enclosing the inflorescence in bud. Inflorescence a cylindric spike, flower without individual bract. Tepals 6, united in lower half. Stamens 6, free. Three outer filaments dilated below and with side teeth. Ovary with 2 ovules in each cell, style ± gynobasic, young ovary with the style in a depression at the summit. 2n = 16. Only one sp., Himalayas and Tibet.

4. Nothoscordum Kunth Fig. 29A,D

Nothoscordum Kunth, Enum. 4: 457 (1843), nom. cons. Zoellnerallium Crosa (1975).

Alliaceous odour absent. Leaves linear or filiform. Scape erect after anthesis. Spathe bracts 2 or in N. fictile Macbr. and N. sessile (R.E. Fries) Beau-verd several. Inflorescence with 1-many flowers. Pedicels slender, often of unequal length. Tepals 6, united below, forming a short tube. Stamens 6, inserted in the tepal tube, sometimes united with each other below. Four to 12 ovules in each cell. In = 10,16, 18, 19, 24, 26. Twenty four to 26 spp., Argentina, Central Chile, Bolivia, S Peru, Paraguay, Uruguay and S Brazil. N. bivalve (L.) Britton extending to Canada, and N. borbonicum Kunth (= N. gracile auct. = N. in-odorum auct. = N. fragrans auct.) is a nearly cosmopolitan weed. Allium siculum Ucria and A. tripedale Trautv. from Asia have previously been referred to Nothoscordum. N. fictile Macbr. from Peru and N. sessile (R.E. Fries) Beauverd from NW Argentina deviate from all other Alliaceae in S America by having the pedicels subtended by individual bracts and by apiculate outer tepals. They may belong to an undescribed genus (Guaglianone 1972). N. andinum (Poepp.) Fuentes may be related; it sometimes has individual bracts (Guaglianone 1973) and was placed by Crosa (1975) into a monotypic genus Zoellnerallium, based on cytological and morphological characters.

5. Ipheion Raf.

Ipheion Raf., Fl. Tellur. 2:12 (1837). Beauverdia Herter (1943).

Alliaceous odour in I. uniflorum (Lindl.) Raf., absent in the other species. Scape curving after anthesis. Scape bracts 2, united at one side for about 1/2 their length, at the other for more. Flower solitary, rarely 2, with tepals united for 1/3-2/3. Anthers free from each other, inserted in the tepal tube at 2 levels. 2n = 12 and 20. Three spp., Central Chile, Argentina, Uruguay, S Brazil.

Traub et al. (1955) included Tristagma and some Nothoscordum in Ipheion, but Traub (1963a) changed the name of this taxon to Tristagma. Guaglianone (1972) included some Nothoscordum with solitary flowers into Ipheion but separated Tristagma.

6. Tristagma Poeppig

Tristagma Poeppig, Fragm. Syn. Phan. 8 (1833). Steinmannia Philippi (1884). Garaventia Looser (1945).

Some species with alliaceous odour. Leaves usually linear and flat, rarely canaliculate, terete or filiform. Scape slender, erect after anthesis. Inflorescence with 1-8 [?] flowers. Bracts united below for less than 15% at any margin. Tepals 6, united below. Anthers 6, placed in 2 series in the tube, versatile or basifixed, filaments long to absent. Several ovules in each locule. 2n = 8,16 and 24. Fourteen spp., Chile and Argentina.

7. Schickendantziella (Speg.) Speg.

Schickendantziella (Speg.) Speg., Anal. Mus. Buenos Aires III, 2: 8 (1903).

Schickendantzia Speg. (1896) non Pax (1889).

One to 2 flowers, violet-purple, nodding. Tepals 3, caudate, almost free. Corona absent. Stamens 6, fertile, filaments united below and enveloping the ovary. Stigma thick. Many ovules in each cell. One sp., Argentina (Tucuman).

8. Speea Loes.

Speea Loes., Notizbl. Bot. Gard. Berlin-Dahlem 10:63 (1927). Geanthus (1884), non Raf. (1814).

Spathe bract 1 or 2 separate, the outer with margins united below. Flower 1-3. Tepals 6, free, caudate. Stamens 6, fertile, filaments free or united, anthers basifixed. Two spp., Chile.

9. Leucocoryne Lindl.

Leucocoryne Lindl., Bot. Reg.: t. 1293 (1830); Zoellner, An.

Mus. Hist. Nat. Valparaiso 5: 9-83 (1972), rev. latace Philippi (1889). ? Erinna Philippi (1864). Stemmatium Philippi (1873). Chrysocoryne Zoellner (non Endl.) (1973). Pabellonia Quezada 8t Martic. (1976).

Some species with alliaceous odour. Leaves linear. Scape slender. Spathe bracts linear-lanceolate. Inflorescence with 1-12 flowers. The 6 tepals united below and forming a tube. Stamens 3 or 6, inserted in the tepal tube; when 3, then opposite the outer tepals. Corona of 3 or 6 lobes. Seeds many in each cell. 2n = 12, 18,24. Fifteen to 20 spp., Chile.

  1. volkmannii (Philippi) Traub has been placed in the monotypic genus Latace, having segments of tepals longer than tube and the 3 functional stamens exserted from the tube. Those with 6 functional stamens are named Stemmatium by Philippi and Chrysocoryne by Zoellner (non Endl.) and Pabellonnia by Quesada et al. Ravenna (1978) includes them in Leucocoryne. Erinna Phil, is dubious or extinct. Only one plant, the type, is known, collected near Santiago de Chile. According to the description, like Leucocoryne, but with smaller flowers and a short tepal tube.
  2. Trichlora Baker

Trichlora Baker, Hook. Icon. PI. 1.1237 (1877).

Bulb narrow, tunicated. Leaves 3-4, linear. Scape slender. Spathe bract 2, free from each other. Inflorescence with about 5 flowers. Outer tepals green, lanceolate, free; inner short, scalelike. Stamens 3 fertile, staminodes 3. Corona absent. Stigma with 3 hornlike branches. Two spp., Peru.

11. Solaria Philippi

Solaria Philippi, Linnaea 29: 72 (1858). Gethyum Philippi (1873). Ancrumia Harvey ex Baker (1877).

Leaves 1-2. Scape very short. Spathe bracts 2, linear. Inflorescence of 3-15 flowers. Tepals 6, green, united below. Corona absent or present as small scales. Functional stamens 2 or 3, staminodes 0 or 3 small, united below and with the tepal tube. Stigma undivided. Ovules 2 in each cell. x = 7. Five spp., S Chile and S Argentina.

12. Miersia Lindl.

Miersia Lindley Bot. Reg.: under t. 992 (1826).

Leaves 3-5, linear. Inflorescence with 3-7 flowers. Tepals 6, free, acuminate. Corona of 6 narrow scales outside the oblique urn-shaped staminal cup bearing 6 small anthers. Style short. Stigma small disk-like. Capsule truncate. 2 n = 20 and 21. Three spp., Chile.

13. Gilliesia Lindley

Leaves usually 2, slender towards the base. Scape slender, longer than the leaves. Spathe bracts unequal. Tepals 6, free, rarely 5. Staminal cup slit down one side, functional stamens 3, staminodes

3. Style slightly trilobed. Alliaceous odour recorded. Five spp., Chile.

Selected Bibliography

Crosa, 0. 1975. Las especies unifloras del género Nothos-cordum Kunth y el género Ipheion Rafinesque de la tribu Allieae (Liliaceae). Darwiniana 19: 335-344.

Crosa, 0.1981. Los chromosomas de cinco especies del género Tristagma (Liliaceae). Darwiniana 23: 361-366.

Dahlgren, R.M.T. et al. 1985. See general references.

Daumann, E. 1970. Das Blütennektarium der Monocoty-ledonen unter besonderer Berücksichtigung seiner systematischen und phylogenetischen Bedeutung. Feddes Repert. Spec. Nov. Regni Veg. 80:463-590.

Davis, G.L. 1966. See general references.

Di Fulvio, E. 1973. Sobre elgineceo de Allium y Nothoscordum. Kurtziana 7: 241-253.

Fay, M.F., Chase, M.W. 1996. Resurrection of Themidaceae for the Brodiaea alliance, and recircumscription of Alliaceae, Amaryllidaceae and Agapanthoideae. Taxon 45: 441-451.

Frölich, D., Barthlott, W. 1988. See general references.

Guaglianone, E.R. 1972. Sinopsis de las especies de Ipheion Raf. y Nothoscordum Kunth (Liliáceas) de Entre Rios y regiones vecinas. Darwiniana 17:159-242.

Guaglianone, E.R. 1973. Nothoscordum andinum, especie de Liliaceae nueva para la flora argentina. Darwiniana 18: 3136.

Hegnauer, R. 1963, 1986. See general references.

Hutchinson, F. 1934. The families of flowering plants. Vol. 2, Monocotyledons. 2nd edn. London: MacMillan.

Jaccard, P., Frey, A. 1928. Kristallhabitus und Ausbildungsformen des Calciumoxalats als Artmerkmal. Vier-teljahrschr. Naturforsch. Ges. Zür. 73, Beiheft: 126-161.

Komarov, V.L. 1968. Flora of the USSR. 4 (Russ. edn. 1935). Jerusalem: Israel Progr. Sc. TransL

Krause, K. 1930. Liliaceae. In; Engler, A., Prantl, K. (eds.) Die natürlichen Pflanzenfamilien, 2nd edn. 15a: pp. 249-329. Leipzig: W. Engelmann.

Mann, L.K. 1960. Bulb organization in Allium: some species of the section Molium. Am. J. Bot. 47: 765-771.

Moore, H.E., Jr. 1954-55. The cultivated Allium. Baileya 2: 103-113, 117-124. 3: 137-149,156-167.

Nuñez, O., Fraissinet, N., Rodríguez, R.H., Jones, K. 1974. Cytogenetic studies in the genus Nothoscordum Kunth. I The N, inodorum polyploid complex. Caryologia 27:403-441.

özhatay, N. 1978. The chromosomes of Milula spicata (Liliaceae) Kew Bull. 32:453-454.

Ravenna, P. 1978. Studies in Allieae II. Plant Life 1978: 130151.

Schnarf, K. 1931. Vergleichende Embryologie der Angiospermen. Berlin: Borntraeger.

Schnarf, K. 1948. Der Umfang der Lilioideae im natürlichen System. Österr. Bot. Z. 95: 257-269.

Schulze, W. 1980. Beiträge zur Taxonomie der Liliifloren V. Alliaceae. Wiss. Ztschr. Friedrich-Schiller-Univ. Jena, Math.-Nat. R. 29: 595-606.

Sterling, C., Huang, S. 1972, Notes on the lactifers of Allium, Cáloscordum, Nothoscordum, Tristagma and Tulbaghia. Plant Life 28: 43-46.

Takhtajan, A.L. 1982. See general references.

Traub, H.P. 1963. Tristagma Poepp. Plant Life 19: 60-61

Traub, H.P., Moldenke, H.N. 1955. The genus Ipheion, diagnosis, key to species and synonymy. Plant Life 11: 125-130.

Vosa, C.G. 1975. The cytotaxonomy of the genus Tulbaghia.

Ann. Bot. (Rome) 34:47-121. Wunderlich, R. 1954. Über das Antherentapetum mit besonderer Berücksichtigung seiner Kernzahl, österr. Bot. Z. 101:1-63.

Wunderlich, R. 1959. Zur Frage der Phylogenie der Endos-permtypen bei den Angiospermen, österr. Bot. Z. 106:203293.

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