Liliaceae

The Liliáceas (Liliaceae Juss.) is a family of perennial plants, grasses, bulbous, frequently pertaining to the Liliales order of the monocotiledóneas. It owns near 16 sorts and 600 species. The Liliáceas can be recognized by their quite great flowers with perigonio formed by 6 free tépalos, frequently colored and with extrorsos spots, 6 estambres and a súpero ovary, to tricarpelar and to trilocular. They are widely distributed by everybody, mainly in regions temperate of the Northern hemisphere.

Historical backgrounds

The Liliaceae family was conceived by Antoine Laurent de Jussieu in 1789 and its definition was very ample and artificial: all the species of plants with 6 tépalos and gineceo of súpero ovary were including in this family. In a while it got to include near 300 sorts and 4,500 species and were included within the great Liliales order (Cronquist 1981, Thorne 1992) like the petaloideas monocotiledóneas, a “group” characterized by flowers with showy tépalos and without starch in endosperma. Cronquist (1981) located to the majority of the petaloideas monocotiledóneas with 6 flowers of estambres in a very ample (and now we know that widely polifilético) Liliaceae. Whereas, in other processings, of they have divided the petaloideas monocotiledóneas with 6 estambres in Liliaceae, including species with a súpero ovary, and Amaryllidaceae, that he includes to the species with ínfero ovary (Lawrence 1951). However, with time he got himself to reconnoiter that, defined of that form, the Liliaceae family included a vast and heterogenous repertoire of sorts that were not related filogenéticamente. Several proposals existed to separate small groups of sorts in more homogenous families, but no of them widely was accepted. In the decade of 1980, in the context of a general revision of the classification of the angiospermaes, the Liliáceas was put under a more intense inspection of election returns. Towards that decade ends, the Botanical Gardens of Kew, the British Museum of Natural Sciences and the Botanical Gardens of Edinburgh formed a committee to examine the possibility of separating to the Liliáceas in more homogenous sub-groups, at least for the organization of their herbal ones. The committee recommended that 24 new families instead of the old man are used and widely defined Liliaceae. In last the two decades the studies of DNA and the morphologic data (particularly those related to the reproductive morphology) added to the cladísticos analyzes, have allowed to conclude that the “petaloideas monocotiledóneas” in fact misna does not belong to one botanical family but they are distributed in two different orders: Asparagales and Liliales. Monofilia of both orders is sustained by cladísticos analyzes based on morphology, ADNr 18S, and many other ADN sequences.

Such studies, added to the analyzes realized within each one of the orders have allowed to regroup previously to the sorts including within Liliaceae in an enormous amount of families, as it is next through cladogramas for the Asparagales and Liliales (according to Judd ET. 2007, modified of Soltis ET to. 2005):

Both cladogramas show that the classic definition of Liliaceae was incorrect, extremely artificial and it did not reflect the true filogenéticas reports between the species composed that it. Therefore, and in spite of the extended use of the ample circumscription of the Liliáceas, the botanists worldwide were adopting the strictest definition of this family and the segregation of the sorts that two orders and numerous families composed in.

Description

Perennial grass with bulbs or rizomas, simple hairs. The roots are typically contráctiles.

The leaves are alternating and espiraladas (and even verticiladas, as in Lilium and Fritillaria) and they are stipulated throughout the stem or in a basal rosette. They are simple, whole, with parallel venación (although in Prosartes and Tricyrtis with clearly reticulada venación between the primary veins), often are envainadoras in the base. Rarely the leaves are pecioladas. They do not appear estípulas.

The inflorescencia usually is determined, sometimes reduced to a unique flower, and terminal. When it is multiflora the flowers are stipulated rarely in a cluster or in one umbela. The flowers are hermafroditas, actinomorphic or zigomorfas, great and slightly generally showy, pediceladas, can or not appear brácteas. Perigonio is formed by 6 tépalos stipulated in two trímeros whirls, are to each other separated and free of the other floral pieces, overlapped, they are petaloideos and usually they appear spots, points or lines of other colors or tones. Perianto can be homoclamídeo (that is, all the tépalos are equal to each other, as in Fritillaria) or diclamídeo (both whirls of tépalos appear morphologic differences, like in Calochortus). The nectar takes place in nectarios in the base of the tépalos.

The androceo appears 6 estambres stipulated in 2 also trímeros whirls, the filaments are to each other separated and free of the other floral pieces. Characteristically, the androceo is diplostémono (that is to say that the external whirl of estambres is opposed to the external tépalos and the internal cycle it is opposed the tépalos interns). The anthers are united to the filament in peltada form or pseudo-basifijas (the end of the filament surrounded but not adhered to the connective tissue), and of longitudinal dehiscence. Pollen is generally monosulcado.

The gineceo is of súpero ovary and is formed by 3 connados carpels, is to trilocular. It appears a single style and a 3-lobado stigma or, 3 stigmata more or less elongados that extend throughout the expensive intern of the branches of the style. The eggs are numerous, with axillary placentación, usually with a tegumento and megasporangio more or less thin. The embryonic coat (megagametofito or gametofito feminine) is variable according to the considered sort. It can be monospórico (Polygonum type) or tetraspórico (Fritillaria type).

The fruit is a loculicida or septicida cap, occasionally a berry. The seeds are flat and with form of disc or globosas, the tegumento is not black, characteristic that them difference of other families, like for example Alliaceae. Endosperma is oily (with aleurone and oils) but without starch, its airframes can be triploides or pentaploides, following the considered embryonic coat.

From a fitoquímico point of view, the family appears esteroideas saponinas but she exhibits crystals of rafidio neither acid chelidónico nor sulphured compounds derived from cisteína (that is, they do not have the scent “aliáceo” characteristic of garlic and the onions). Basic the chromosomal numbers are variable according to the considered sort.

To see a recent processing of the family in Tamura (1998b).

Ecology

The Liliáceas widely is distributed, mainly in regions temperate of the Northern hemisphere. Generally they are plants of level prairies, mountain meadows and other open communities. They own its center of diversity in the southwest from Asia to China.

Usually they bloom in the primrose. The showy flowers of this family are polinizadas by insects, especially bees, wasps, butterflies, and polillas. The nectar or pollen that produces the flowers of the liliáceas in enormous amount is used like reward of the pollination. The seeds are dispersed so much by the wind as by the water, a few species have structures of type seed coat and are dispersed by ants.

Philogeny

Liliaceae, in its strict current definition, is clearly monofilética (Chase ET to. 1995a, b), although it is difficult to diagnose morphologically (Tamura 1998b). Hayashi and Kawano (2000) secuenciaron the regions rbcL and matK of the cloroplastídico DNA of Lilium and related sorts and found coherent results with the circumscription of propose Liliaceae sensu stricto by Tamura (1998, that separated the subfamily Calochortoideae in its own Calochortaceae family). Patterson and Givnish (2002) secuenciaron the genes rbcL and ndhF and found a fort you maintain for monofilia of Liliaceae, the publication also describes a philogeny of the sorts and provides estimations with the times of divergence.

Calochortus, Prosartes, Scoliopus, Streptopus, and Tricyrtis forms clado (the subfamily Calochortoideae), and is grass with rizomas crawling (“creeping”), styles divided in the apex, and the development of the megagametofito of the Polygonum type (that is to say, the megagametofito being developed of a simple megaspora, with endosperma triploide). In other Calochortus classifications it has been located in his own family, whereas the other sorts of Calochortoideae were located in a heterogenous Uvulariaceae (Dahlgren ET to. 1985) or in an expanded Calochortaceae (Tamura 1998a). Nevertheless, these members of Liliaceae nearly are not related to the morphologically similar Uvularia and Disporum (Shinwari ET to. 1994), and the last one here is located in Colchicaceae.

The rest of the sorts of Liliaceae constitutes the Lilioideae, great clado characterized by bulbs and contráctiles roots, and a megagemetofito being developed of 4 megasporas (of Fritillaria type).

Monofilia of each of these two subclados of Liliaceae is maintained by characters of ADN sequences (Chase ET to. 1995a, 2000, Patterson and Givnish 2002).

Third clado was identified from the filogenéticas investigations of Clintonia, a sort of distribution disyunta between North America and the east of Asia. Using the 5 species of Clintonia (andrewsiana Clintonia, Clintonia borealis, umbellulata Clintonia, uniflora Clintonia and Clintonia udensis), several representatives of Liliaceae (such as Cardiocrinum cordatum, virginiana Medeola, Scoliopus bigelovii and Scoliopus hallii) and the data of the sequences of the cloroplastídicos genes rbcL and matK, reached the conclusion that Clintona is monofilético, that consists of two clados, one of Eastern Asia and the other of North America, and that the sort more closely related is Medeola. Both sorts, Clintonia and Medeola, were stipulated in a separated subfamily, Medeoloideae.

Taxonomy

The subfamilies of Liliaceae

• 1. Lilioideae Eaton (synonymous: Erythroniaceae Martynov, Fritillariaceae R.A.Salisbury, Liriaceae Batsch, Medeolaceae Takhtajan, Tulipaceae Batsch). It frequently includes bulbous plants and with contráctiles roots. The stems are not graft, the leaves have parallel nerviación. The flowers are great and the embryonic coat is of the Fritillaria type (that is, tetraspórico). The fruit is a septicida cap. The seeds frequently are flattened, with the exotesta in lignificada fence or. Basic the chromosomal number is variable (n=9, 11 to 14). The chromosomes, as well, appear a very variable longitude, that goes from the 2.2 µm to the 27 µm. This subfamily understands 11 sorts and around 535 species that distribute in regions temperate and cold of North America and the east of Asia. The sorts of Lilioideae with greater wealth of species are Tulipa (150 species), Fritillaria (130), Lilium (110) and Gagea (90).

• 2. Calochortoideae Dumortier. They appear great leaves with parallel nerviación, flowers, without style or with a very short style. They own perianto differentiated in chalice and corola, with the pubescentes tépalos. The fruit is a cap with septicida dehiscence and the embryonic coat is of the Polygonum type. The chromosomal numbers vary of n=6 to n=13, with the 6.5 chromosomes from 1.5 to µm of longitude. It includes/understands 5 sorts, Calochortus, Prosartes, Scoliopus, Streptopus and Tricyrtis, and 100 species distributed in the east of North America and the east of Asia. Several taxonomists have suggested these 5 sorts must be secreted of Liliaceae and to stipulate themselves in their own family, Calochortaceae. Despite the previous thing, the modern systems of classification (as APG or APG II) do not reconnoiter this last family and include to Calochortus and compatible sorts within the Liliáceas.
• 3. Medeoloideae. It includes 2 sorts (Clintonia and Medeola) and 7 species distributed in the east of Asia and North America. They appear seeds with striae and basic a chromosomal number of n=7.

Listing of sorts

As it were described in the historical backgrounds, the definition of Liliaceae has changed in the course of the years. In the past one treated to the family like a great heterogenous assembly (broad Liliaceae sensu) that, more recently, it was divided in numerous secreted families. In the modern circumscription of Liliaceae one considers that the family is composed by 16 sorts and 635 species, which list themselves next:

• Calochortus Pursh (incl.: Cyclobothra Sweet).
• Cardiocrinum (Endl.) Lindl.
• Clintonia Raf.
• Erythronium L.
• Fritillaria L. (incl.: Orithyia D. Don, Petilium Ludw., Rhinopetalum Fisch. ex- D.Don, Korolkowia Regely Theresia K.Koch).
• Gagea Salisb.
• Lilium L.
• Lloydia Salisb. ex- Rchb. (incl.: Giraldiella Dammer).
• Medeola L.
• Nomocharis Franch.
• Notholirion Wall. ex- Boiss.
• Prosartes D. Don (without.: Disporum Salisb.).
• Scoliopus Torr.
• Streptopus Michx.
• Tricyrtis Wall. (incl.: Brachycyrtis Koidz., Compsoa D. Don).
• Tulipa L. (incl.: Deep Amana).

The represented sorts more are Fritillaria (100 species), Gagea (90 species), Tulipa (80 species), Lilium (80 species), and Calochortus (65 species).

Economic importance

It is also seen: Bulbous plants, Flower-growing

Many species of the Tulipa sorts (“tulipán”), Fritillaria (“ajedrezada”), Lilium (“iris”, “lily”), Calochortus and Erythronium are cultivated like ornamental plants anywhere in the world. However, the tulips and the lilies or irises are those of greater economic importance within the family, as it is described next.

Tulipán

Tulipán is cultivated with two main objectives, the production of cut flower and the one of bulbs droughts. These last ones are destined, as well, to satisfy the demand of bulbs for parks, gardens and family use and, on the other hand, to provide necessary bulbs for the production with cut flower. The international trade of cut flower has an approximated overall value of 11,000 million Euros, which provides a magnitude with the economic importance of this activity.

The main producing bulb country of tulipán is Holland, country that concentrates 87% of the cultivated world-wide area, which is of approximately 12,000 hectares. The bulbs of this species take place significantly in other 14 countries, headed by Japan, France and Poland. The majority of these countries uses bulbs obtained for its own production of cut flower or to supply to its retail bulb market droughts. Holland, nevertheless, aside from being the main international bulb producer, is the exception to this generalization. In fact, it annually produces approximately 4,000 million bulbs, of which 53% are used in the market of cut flower and the rest are used in the bulb market droughts. Of bulbs destined to the market of cut flower, Holland uses 57% to satisfy its internal market and it exports it to the rest to several countries, inside and outside the European Union.

Irises and lilies

Of similar way that in the case of the tulips, the greater area of production of the varieties of Lilium is concentrated and Holland (76% of world-wide area). Other 9 countries, headed by France, Chile, Japan, Member States and New Zealand, produce lilies on significant scale. Half of the producing countries uses the bulbs that produce in the industry of flower-growing and only one small proportion destines to the market of bulb sales droughts. Countries such as Holland, France, Chile, New Zealand and Australia use produced bulbs to supply not only their internal market but also to export them. Holland produces annually around 2,200 million iris bulbs, of which 96% are used for their own production of flowers and the rest is exported to countries of the European Union mainly.

Notes

a. ↑ the embryonic coats can be of different types according to how many megasporas they degenerate after the meiosis that undergoes the airframe mother of the megaspora. In the Polygonum type they degenerate three of the 4 megasporas and for that reason the embryonic coat says monospórico (espora comes from a single). In the embryonic coat of Fritillaria type no of the 4 megasporas degenerates after the meiosis and all of them participate in the structure of the mature embryonic coat, for that reason “tetraspórico” is denominated.

Mentioned references

1. ↑ Jussieu, A.L.. (1789), It generates Plantarum, 48.
2. ↑ PlantSystematics.org. Consulted 2008-05-24.
3. ↑ Stevens, P.F. (2001 in future). Liliaceae (in English). Angiosperm Phylogeny Website. Version 7, May 2006. Consulted 2008-28-04.
4. ↑ Cronquist, A. (1981), An integrated system of classification of flowering plants., New York: Columbia University Press.
5. ↑ Thorne, R.F. (1992). “Classification and geography of the flowering plants.”. Bot. Rev. (58): 225-348. Consulted the 25/02/2008.
6. ↑ Lawrence, G.H.M. (1951), vascular Taxonomy of plants., Macmillan.
7. ↑ Mathew, Brian (1989), Splitting the Liliaceae, The Plantsman 11 (2): 89.
8. ↑ Chase, M.W.; Duvall, M.R., Hills, H.G., Conran, J.G., Cox, A.V., Eguiarte, L.E., Hartwell, J., Fay, M.F., Caddick, L.R., Cameron, K.M., and Hoot, S. (1995), “Molecular systematics of Lilianae.” Rudall, P.J., Cribb, P.J., Cutler, D.F., Monocotyledons: Systematics and evolution., Royal Botanical gardens, 109-137.
9. ↑ Chase, M.W.; Stevenson, D.W., Wilkin, P., and Rudall, P.J. (1995b), “Monocot systematics: To combined analysis.” Rudall, P.J., Cribb, P.J., Cutler, D.F., Monocotyledons: Systematics and evolution., Royal Botanical gardens, 685-730.
10. ↑ Chase, M.W.; Soltis, D.E., Soltis, P.S., Rudall, P.J., Fay, M.F., Hahn, W.H., Sullivan, S., Joseph, J., Molvray, M., Kores, P.J., Givnish, T.J., Sytsma, K.J., and Pires, J.C. (2000), “Higher-level systematics of the monocotyledons: An assessment of current knowledge and to new classification.” Wilson, K.L. and Morrison, D.A., Monocots: Systematics and evolution., CSIRO Publ., 3-16.
11. ↑ Chase, M.W., Fay, M.F.; Devey, D.S.; Maurin, Or; Rønsted, N; Davies, T.J; Pillon, and; Petersen, G; Seberg, Or; Tamura, M.N.; Lange, Conny Bruun Asmussen (Faggruppe Botanik); Hilu, K; Borsch, T; Davis, J.I; Stevenson, D.W.; Pires, J.C.; Givnish, T.J.; Sytsma, K.J.; McPherson, M.A.; Graham, S.W.; RAI, H.S. (2006). “Multigene analyzes of monocot relationships: to summary” (pdf). Alder (22): 63-75. ISSN: 00656275. Consulted the 25/02/2008.
12. ↑ Davis, J.I., Stevenson, D.W.; Petersen, G.; Seberg, Or.; Campbell, L.M.; Freudenstein, J.V.; Goldman, D.H.; Hardy, C.R.; Michelangeli, F.A.; Simmons, M.P.; Specht, C.D.; Vergara-Silva, F.; Gandolfo, M. (2004). “To phylogeny of the monocots, for ace inferred from rbcL and atpA sequence variation, and to comparison of methods calculating jacknife and bootstrap VALUEs.”. Syst. Bot. (29): 467-510. Consulted the 25/02/2008.
13. ↑ Graham, S.W., Zgurski, J.M., McPherson, M.A., Cherniawsky, D.M., Saarela, J.M., Horne, E.S.C., Smith, S. and., Wong, W.A., O'Brien, H.E., Biron, V.L., Pires, J.C., Olmstead, R.G., Chase, M.W., and RAI, H.S. (2006). “Robust inference of monocot deep phylogeny using an expanded multigene plastid dates Seth.” (pdf). Alder (22): 3-21. Consulted the 25/02/2008.
14. ↑ Hilu, K., Borsch, T., Muller, K., Soltis, D.E., Soltis, P.S., Savolainen, V., Chase, M.W., Powell, M.P., Stakes out, L.A., Evans, R., Sauquet, H., Neinhuis, C., Slotta, T.A.B., Rohwer, J.G., Campbell, C.S., and Chatrou, L.W. (2003). “Angiosperm phylogeny based on matK sequence information.”. American J. Bot. (90): 1758-1766. Consulted the 25/02/2008.
15. ↑ Pires, J.C., Maureira, I.J., Givnish, T.J., Sytsma, K.J., Seberg, Or., Petersen, G., Davis, J.I., Stevenson, D.W., Rudall, P.J., Fay, M.F., and Chase, M.W. (2006). “Phylogeny, genome size, and chromosome evolution of Asparagales.”. Alder (22): 278-304. Consulted the 25/02/2008.
16. ↑ Rudall, P., Furness, a.c., Chase, M.W., and Fay, M.F. (1997a). “Microsporogenesis and pollen sulcus type in Asparagales (Lilianae).”. Canad. J. Bot. (75): 408-430. Consulted the 25/02/2008.
17. ↑ Källersjö M, JS Farris, MW Chase, B Bremer, MF Fay, CJ Humphries, G Petersen, Or Seberg, and K Bremer (1998). “Simultaneous parsimony jacknife 2538 analysis of rbcL DNA sequences reveals support for major clades of green plants, land plants, and flowering plants.”. Pl. Syst. Evol. (213): 259-287. Consulted the 25/02/2008.
18. ↑ Fay, M.F. (2000), “Phylogenetic studies of Asparagales based on four plastid DNA regions.” K.L. Wilson and D.A. Morrison, Monocots: Systematics and evolution., Royal Botanical gardens, 360-371, Kollingwood, Australia: CSIRO.
19. ↑ Soltis OF, PS Soltis, MW Chase, ME Mort, DC Albach, M Zanis, V Savolainen, WH Hahn, SB Hoot, MF Fay, M Axtell, SM Swensen, LM Prince, WJ Kress, KC Nixon, and JS Farris. (2000). “Angiosperm phylogeny inferred from 18S rDNA, rbcL, and atpB sequences.”. Bot. J. Linn. Soc. (133): 381-461. Consulted the 25/02/2008.
20. ↑ Stevenson, D.W.; Davis, J.I., Freudenstein, J.V., Hardy, C.R., Simmons, M.P., and Specht, C.D. (2000), “To phylogenetic morphological analysis of the monocotyledons based on and molecular character sets, with comments on the placement of Acorus and Hydatellaceae.” Wilson, K.L. and Morrison, D.A., Monocots: Systematics and evolution., CSIRO Publ., 17-24.
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22. ↑ Goldblatt, P. (1995), “The status of R. Dahlgren's orders Lliliales and Melanthiales.” Rudall, P.J., Cribb, P.J., Cutler, D.F., and Humphries, C.J., Monocotyledons: Systematics and evolution., Royal Botanical gardens, 181-200.
23. ↑ Stevenson, D.W.; Loconte, H. (1995), “Cladistic analysis of monocot families.” Rudall, P.J., Cribb, P.J., Cutler, D.F., Monocotyledons: Systematics and evolution., Royal Botanical gardens, 543-578.
24. ↑ Vinnersten, A., Bremer, K. (2001). “Major Age and biogeography of clades in Liliales”. Amer. J. Bot. (88): 1695-1703. Consulted the 25/02/2008.
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27. ↑ Hayashi, K., Kawano, S. (2000). “Molecular systematics of Lilium and allied generates (Liliaceae): Phylogenetic relationships among Lilium and related generates based on the rbcL and matK gene sequence dates.”. Plant Species Biology 15:73 - 79. Consulted the 23/03/2008.
28. ↑ Patterson, T.B., Givnish, T.J. (2002). “Phylogeny, concerted convergence, and phylogenetic niche conservatism in the Core Liliales: Insights from rbcL and ndhF sequence dates.” (pdf). Evolution 56:233 - 252. Consulted the 23/03/2008.
29. ↑ Dahlgren, R.M.; Clifford, H.T., Yeo, P.F., The families of the monocotyledons., Springer-Verlag.
30. ↑ Tamura, M.N. (1998), “Calochortaceae” K. Kubitzki, The families and generates of vascular plants, vol. 3, Monocotyledons: Lilianae (except Orchidaceae). , 164-172, Berlin: Springer-Verlag.
31. ↑ Shinwari, Z.K., Terauchi, R., Hteck, F.H., and Kawano, S. (1994). “Recognition of the New World Disporum section Prosartes (Liliaceae) based on the sequence dates of the rbcL gene.”. Taxon (43): 353-366. Consulted the 25/02/2008.
32. ↑ Kazuhiko Hayashi, Seiji Yoshida, Frederick H. Utech, Shoichi Kawano (2001). “Molecular systematics in the genus Clintonia and related taxa based on rbcL and matK gene sequence dates”. Plant Species Biology 16 (2), 119-137 doi: 10.1046/j.1442-1984.2001.00057.x. Consulted the 25/04/2008.
33. ↑ Dahlgreen R.M.T., Clifford H.T. and Yeo P.F., 1985, The families of the monocotyledons: structure, evolution and taxonomy. Springer Verl., Stuttgart
34. ↑ Ownbey, M. 1940. To monograph of the genus Calochortus. Ann. Missouri Bot. Gard. 27:371 - 560.
35. ↑ McDonald, H.P. To review of Calochortus. Herbertia 55:34 - 51; 2000
36. ↑ Patterson, T.B., and Givnish, T.J., 2002, Phylogeny, concerted convergence, and phylogenetic niche conservatism in the Core Liliales: Insights from rbcL and ndhF sequence dates: Evolution, v. 56, P. 233-252.
37. ↑ Genera of Liliaceae (in English). USDA, ARS, National Genetic Resources Program. Germplasm Resources Information Network - (GRIN). National Germplasm Resources Laboratory, Beltsville, Maryland. (2001 in future). Consulted the 2008-4-05.
38. ↑ http://www.aboutflowerbulbs.com/bulb_globalization.htm
39. ↑ Frankel, R. & Galun, E. (1977), Pollination mechanisms, reproduction, and plant breeding, Monographs on Theoretical and Applied Genetics no. 2. Springer-Verlag. Berlin. ISBN 0387079343.

Bibliography

• Judd, W.S.; C.S. Campbell, E.A. Kellogg, P.F. Stevens, M.J. Donoghue (2007), “Liliaceae” Plant Systematics: To Phylogenetic Approach, 3ª editing, Sunderland, Massachusetts: Sinauer Associates. ISBN 9780878934072.

• Simpson, Michael G. (2005), “Liliaceae” Plant Systematics., 180, Inc. Elsevier. ISBN 0126444609 ISBN-13 9780126444605.

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