Liliaceae—i

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Liliaceae—i

Description

Herbs, usually glabrous, with perennial underground stems (corms, bulbs, tubers, or rhizomes) in all Mal. spp. Leaves simple, caespitose and basal, sometimes distichous, if cauline usually alternate, generally linear to lanceolate or oblanceolate especially when basal, but sometimes shorter and broader (to ovate) when cauline, usually sessile (in Asparagus and Petrosavia reduced to non-photosynthetic scales), usually with parallel venation. Stipules 0. Inflorescence terminal or axillary, usually racemose (less often at least partly umbellate) or flowers solitary, usually bracteate. Flowers bisexual (except, in Mal., Asparagus cochinchinensis and Astelia alpina), usually actinomorphic. Stamens 6, inserted on receptacle or perianth; Ovary usually superior, of 3 (usually fused) carpels; Fruit usually a loculicidal or septicidal capsule or berry, rarely the ovary wall ruptured by the developing seed which develops unprotected by a fruit, perianth caducous or persistent. Seeds with copious fleshy or cartilaginous endosperm.

Distribution

Africa: present Asia: present Asia-Tropical:, Malayapresent; New Guineapresent; Philippines (Philippinespresent); Sumatera (Sumaterapresent); Thailand (Thailandpresent) Australasia: present East Malesia: present Himalaya: present Mauritius: present Old World: present Pacific Islands: present S. China: present South Africa: present South America: present all over the world: present alpine zone of New Guinea: present northern hemisphere: present
About 180 genera with approximately 3500spp., distributed all over the world, especially in the temperate regions of Asia, Australasia and Africa, but relatively poorly represented in South America (13 genera).
In Malesia 22 genera, with a total of 31 spp., no genus being represented by more than two species. The only genus endemic to the region is the Malayan genus Tricalistra whose separation from Tupistra is, however, somewhat uncertain. Most other genera are represented in Malesia by a minority of their species, exceptions being Gloriosa and Peliosanthes, which are probably both monotypic, and Petrosavia, which consists probably of two species.
The genera can roughly be arranged into three geographical groups.
Old World genera are Asparagus, Chlorophytum, Dianella, Gloriosa and Iphigenia, among which Chlorophytum is mainly Africa-centred, and Dianella mostly Australasian.
Northern hemisphere genera, especially from the Far East, Sino-Himalayan, are the following: Aletris, Disporopsis*, Disporum, Lilium*, Liriope*, Ophiopogon, Peliosanthes, Petrosavia, Tricyrtis*, and Tupistra. Of these four, provided with an asterisk, are found only in Malesia in the Philippines, and Tupistra only in Malaya and Sumatra. All of them are absent from East Malesia. Most of their species occur in the montane zone, testimony of their subtemperate ecology.
Australasia-derived genera are Arthropodium, Astelia, Caesia, Schelhammera, Thysanotus, and Tricoryne. Their occurrence in Malesia is confined to New Guinea, except for Thysanotus chinensis Bth. which is found through Malesia as far as Thailand and S. China. Their ranges are sometimes wider in Austral regions, as Arthropodium and Caesia occur also in the Malagasy area, and Caesia also in South Africa, while Astelia ranges widely from Mauritius to the southern Pacific islands and the Falkland Islands. Except for Astelia, which occurs in Malesia only in the alpine zone of New Guinea, all the species of the genera of this southern group are bound to lowland drought habitats.

Dispersal

The great majority of Liliaceae spread and reproduce vegetatively by the branching of their subterranean axes. In most species this appears to be a slow process, with the branches often not extending more than a few centimetres in a year. It may, however, result in fairly dense monospecific stands, for example, in Astelia and Liriope.
Fruits are generally capsules or berries. In the former, dispersal mechanisms do not usually result in the removal of seeds to any great distance, although wind and water can contribute significantly.
Birds are probably the most efficient vectors over longer distances. Several genera have fleshy fruits and two (Ophiopogon and Peliosanthes) have a fleshy coating to the seeds, which are exposed through rupture of the ovary wall. Mammals may also disperse the seeds by eating the fruits. Liliaceae seeds in other areas are known to be carried by ants if there is a substance attractive to ants (often oil bodies) in the testa or fruit. Specific data on the Malesian species have not, however, been found.
For a study of the structure and relationships of the seeds, see .

Taxonomy

The family Liliaceae, in the sense of Bentham & Hooker and of Krause in E. & P., Nat. Pfl. Fam. is a very large and rather heterogeneous one including possibly as many as 3500 species. Many more recent authors have attempted to distribute these species over a larger number of families. In this treatment the family delimitation of Hutchinson () has been adopted, with two modifications: the inclusion of Petrosavia which Hutchinson placed in its own family, and of the naturalised Nothoscordum which Hutchinson placed in the Amaryllidaceae. It is very doubtful if the family is more naturally defined by excluding a number of genera represented in Malesia, as Hutchinson has done, but this has been followed here as much for the convenience of dealing with smaller families as for any conviction that these families have any botanical significance. Table 1 indicates the genera retained in the Liliaceae and the families to which other genera, sometimes included in the Liliaceae, were ascribed by Hutchinson. These families have also been included in the key to the genera of Malesian Liliaceae.
Much work remains to be done on the relationships of Liliaceous genera and Hutchinson's work, although the most recent, is probably no better than Krause's. Hutchinson's placing of Ophiopogon and Peliosanthes in separate tribes is, for example, almost certainly unjustified.
Table 1. Malesian Liliaceous genera in the classification by Krause (1930), first column, and by Hutchinson (1973), third column.
Subfamily I.Melanthioideae
Subfamily I.2.PetrosavieaePetrosavia(Petrosaviaceae)
Subfamily I.6.UvularieaeSchelhammeraUvularieae
Subfamily I.6.UvularieaeGloriosaUvularieae
Subfamily I.7.TricyrteaeTricyrtisTricyrtideae
Subfamily I.8.AnguillarieaeIphigeniaIphigenieae
Subfamily III.Asphodeloideae
Subfamily III.11.AsphodeleaeChlorophytumAsphodeleae
Subfamily III.11.AsphodeleaeThysanotusAsphodeleae
Subfamily III.11a. AsphodelinaeArthropodiumAsphodeleae
Subfamily III.11a. AsphodelinaeTricoryneJohnsonieae
Subfamily III.11a. AsphodelinaeCaesiaAsphodeleae
Subfamily III.11g. DianellinaeStypandraDianelleae
Subfamily III.11g. DianellinaeDianellaDianelleae
Subfamily III.17.LomandreaeLomandra(Xanthorrhoeaceae)
Subfamily III.17.LomandreaeRomnalda(Xanthorrhoeaceae)
Subfamily IV.AllioideaeNothoscordum (introduced)(Amaryllidaceae)
Subfamily V.LilioideaeLiliumTulipae
Subfamily VII.Dracaenoideae
Subfamily VII.27.DracaeneaeCordyline(Agavaceae)
Subfamily VII.27.DracaeneaeDracaena(Agavaceae)
Subfamily VII.27.DracaeneaeAsteliaMilliganieae
Subfamily VIII.Asparagoideae
Subfamily VIII.28.AsparageaeAsparagusAsparageae
Subfamily VIII.29.PolygonataeDisporumPolygonatae
Subfamily VIII.29.PolygonataeDisporopsisPolygonatae
Subfamily VIII.30. ConvallarieaeTupistraAspidistreae
Subfamily VIII.30. ConvallarieaeTricalistraAspidistreae
Subfamily IX.MondoideaeLiriopeOphiopogoneae
Subfamily IX.MondoideaeOphiopogonOphiopogoneae
Subfamily IX.MondoideaePeliosanthesPeliosantheae
Subfamily X.AletroideaeAletrisNarthecieae
Subfamily XI.LuzuriagoideaeLuzuriaga(Philesiaceae)
Subfamily XI.LuzuriagoideaeGeitonoplesium(Philesiaceae)
Subfamily XI.LuzuriagoideaeEustrephus(Philesiaceae)
Subfamily XII.SmilacoideaeRhipogonum(Smilacaceae)
Subfamily XII.SmilacoideaeSmilax(Smilacaceae)
Subfamily XII.SmilacoideaeHeterosmilax(Smilacaceae)

Cytology

Liliaceae, because of the usually large size of their chromosomes, because of the ease with which material can often be obtained at the stages of division required for study, and because many species are in cultivation, have been fairly well studied cytologically. Chromosome numbers, and even basic chromosome numbers vary widely sometimes within, as well as between, genera. At least six different somatic numbers have, for example, been reported for Disporum, based on x = 6, 7, 8, 9 and 11. Other genera, for example, Asparagus (x = 10) and Dianella (x = 8), have relatively stable basic numbers, although polyploidy may be common.

Uses

Several species of Liliaceae native to Malesia have been taken into cultivation as garden ornamentals, for example of the genera Dianella, Gloriosa, Lilium, Liriope, and Ophiopogon. Other uses are, however, rather few. Several genera, for example, Asparagus, Gloriosa, Ophiopogon, have been used in traditional medicines but they have not contributed to modern medicine. Others have edible underground parts (e.g. Arthropodium) or fruits (e.g. Astelia), but none is probably of great significance.

Notes

A very large number of exotic Liliaceae are recorded to be or have been cultivated in gardens in Malesia. They have been treated elaborately, with keys for their identification by , in Dutch, and by in English.
Only one exotic, Nothoscordum inodorum, has been introduced and has run wild in West Java; this has been incorporated in the treatment.
Almost all drawings were made by Mr. L. Dutkiewicz, Adelaide.