Amaryllidaceae J.St.-Hil.

Meerow, A. (2009). Neotropical Amaryllidaceae.

Morphology

Description

Habit : bulbous (rarely rhizomatous), mostly geophytic, perennials, terrestrial, occasionally aquatic or epiphytic , rich in family-specific alkaloids; bulbs tunicate.  Leaves annual or persistent , sessile and linear or lorate, or petiolate and lanceolate to widely elliptic , distichous or spirally arranged; sometimes basally sheathing and forming an aerial pseudostem, usually glabrous , rarely with trichomes; phyllotaxy chiefly distichous , sometime spiralled.  Inflorescence scapose, pseudo-umbellate (reduced helicoid cymes); scape sometimes wholly subterranean, appearing obsolete, terminated by two or more spathaceous, obvolute or equitant, usually marcescent bracts that enclose the flowers in bud (bracts rarely absent); inner bracteoles usually present and successively shorter and narrower.  Flowers 1-many, perfect, frequently large and showy, sessile or pedicellate, each usually subtended by a bracteole , actinomorphic or zygomorphic , generally protandrous; perigone crateriform, salverform, funnelform, tubular or ventricose, consisting of 3 + 3 segments (tepals) connate below into a short or long tube or rarely free to the base; inner tepals generally shorter than the outer; outgrowth of the perigone sometimes present, forming a conspicuous corona (paraperigone), or relatively inconspicuous and consisting of a short callose rim or ring of scales or fimbriae at the throat; stamens 3 + 3, rarely 5 or 18 or more, subequal or varying in length, inserted at the perigone throat or below, the filaments sometimes variously connate or otherwise appendaged, rarely adnate to the style ; anthers usually dorsifixed, rarely centrifixed or basifixed, introrse, dehiscing longitudinally or rarely from a terminal pore; style filiform , occasionally strumose, rarely tripartite; stigma capitate , 3- lobed or deeply trifid, usually papillate; ovary syncarpous, tri-carpellate, inferior, tri- (rarely uni-) locular, with septal nectaries; ovules axile or basal in placentation, anatropous, crassinucellate, bi-, uni- or ategmic.  Fruit a loculicidally dehiscent capsule , sometimes indehiscent , rarely baccate ; seeds globose or subglobose and fleshy or hard, or flattened and winged , usually with a black or brown phytomelanous testa, sometimes with a caruncular elaiosome at the chalazal end; endosperm with hemicellulose and lipids, in the more derived genera rich in water and/or starch.

Distribution

Distribution in the Neotropics

• In the Neotropics, the family occurs from Mexico through Central America and the West Indies to Chile and Argentina in South America. Notable areas of diversity throughout this range include eastern Brazil, north-central Chile (outside of the tropical zone, however), and the central Andes of Ecuador and Peru.
• HippeastrumHerb. is primarily found in the Andes and eastern Brazil, Hymenocallis Salisb. occurs mostly in Mesoamerica, ClinanthusHerb. is largely endemic to Peru, and ZephyranthesHerb. is broadly distributed. The greatest generic diversity is found in Peru.
• The Neotropical genera of Amaryllidaceae are chiefly adapted for seasonally dry habitats and some prefer truly xeric environments in which their bulbs may remain dormant for a period longer than they are in active growth (e.g., Leptochiton Sealy, Paramongaia Velarde, some Eucrosia Ker Gawl.). At the other extreme, species have colonized the understory of rain forests (Eucharis Planch. & Linden, Griffinia Ker Gawl.) and aquatic habitats (a number of Hymenocallis, Hippeastrumangustifolium Phil., Crinum L.).
• The family has also adapted to the high montane tropical climates of the Andes. Certain genera are primarily found at elevations in excess of 2,000 meters; and Clinanthushumilis (Herb.) Meerow is found above 4,000 meters. This species has adapted to high elevations by retaining the scape (and developing fruit) inside the bulb until the seeds are ripe.

Diagnostic

Distinguishing characters (always present)

• Bulbous herbs.
• Leaves distichous, rarely spiral, simple; blades mostly linear or strap-shaped, the base often sheathing a short stem, sometimes pseudopetiolate.
• Inflorescences umbels, borne on scapes.
• Flowers with 6 tepals in 2 whorls, usually similar in shape and color; stamens 6; ovary inferior.
• Fruits loculicidal capsules.

Key differences from similar families

• From Alliaceaes.s.: lack of allyl sulfide compounds, inferior ovary.
• From Agapanthaceae: lack of steroid saponins, inferior ovary.
• From Alliaceaes.s. and Agapanthaceae: presence of unique amaryllidaceous alkaloids.

Other important characters

• Seed testa with phytomelan crust.

General Description

Status

• All of the genera listed except Crinum are endemic to the Americas.
• In addition to the native species, there are several African Crinum species that are naturalized in the Neotropics.
• Various other exotic members of the family are cultivated in the Neotropics, as are hybrids of native genera such as Hippeastrum.

Number of genera

In tropical America, there are 26 genera and about 375 species. The largest genera in the Neotropics are Hippeastrum (50-60 species), Hymenocallis (about 50), Zephyranthes (about 50), and Clinanthus (about 30).

NOTE: The genera Phycella Lindl., Rhodophiala C.Presl and Placea Miers have been excluded as they occur only in Mediterranean and temperate Chile and warm temperate Argentina.

• CaliphruriaHerb.
• ChlidanthusHerb.
• ClinanthusHerb.
• Crinum L.
• Eithea Ravenna
• Eucharis Planch. & Linden
• Eucrosia Ker Gawl.
• Eustephia Cav.
• Griffinia Ker Gawl.
• HabranthusHerb.
• Hieronymiella Pax.
• HippeastrumHerb.
• Hymenocallis Salisb.
• Ismene Salisb.
• Leptochiton Sealy
• Pamianthe Stapf
• Paramongaia Velarde
• PhaedranassaHerb.
• Plagiolirion Baker
• PyrolirionHerb.
• Rauhia Traub
• Sprekelia Heist.
• StenomessonHerb.
• Urceolina Rchb.
• Worsleya Traub
• ZephyranthesHerb.

Notes on delimitation

• Multiple DNA sequences indicate that Amaryllidaceae, Alliaceae and Agapanthaceae form a monophyletic group which can be treated as one family. 
• The AGP II listed both classifications as options. 
• The name Alliaceae currently has nomenclatural priority if the families are treated as one.
• A proposal to super-conserve the name Amaryllidaceae has been submitted to the nomenclatural committee of the International Association for Plant Taxonomy.

Literature

Important literature

Baker, J.G. 1888. Handbook of the Amaryllideae. George Bell and Sons, London.

Herbert, W. 1837. Amaryllidaceae. J. Ridgeway and Sons, London.

Meerow, A.W. and D.A. Snijman. 1998. Amaryllidaceae. In: K. Kubitzki (ed.),  The Families and Genera of Vascular Plants 3: 83-110. Springer-Verlag, Berlin.

Meerow, A.W., M.F. Fay, C.L. Guy, Q-B. Li, F.Q. Zaman, & M.W. Chase. 1999.  Systematics of Amaryllidaceae based on cladistic analysis of plastid rbcL and trnL-F sequence data.  Amer. J. Bot. 86: 1325-45.

Meerow, AW., C.L. Guy, Q. Li, & S-L. Yang.  2000. Phylogeny of the American Amaryllidaceae based on nrDNA ITS sequences.  Syst. Bot. 25: 708-26.

Traub, H.P. 1963. Genera of the Amaryllidaceae. American Plant Life Society, La Jolla, CA.

Amaryllidaceae, Inger Nordal (University of Oslo). Flora of Tropical East Africa. 1982

Morphology General Habit

Herbs with bulbs or rhizomes

Morphology Leaves

Leaves in a basal rosette from the apex of the bulb or rhizome; petioles, if present, sometimes sheathing to form a false stem; blade simple, entire, linear to lanceolate or strap-shaped

Morphology Reproductive morphology Inflorescences Scape

Scape leafless, central or lateral in relation to the leaves, with 1–many flowers in an umbel-like inflorescence, subtended by an involucre of 1–many bracts and with ephemeral hyaline bracts between the flowers

Morphology Reproductive morphology Flowers

Flowers showy, bisexual, 3-merous, regular or less often slightly irregular

Morphology Reproductive morphology Flowers Perianth

Tepals in 2 series, equal to subequal, inserted above the ovary, free or partly united into a short or long tube; corona sometimes present

Morphology Reproductive morphology Flowers Androecium

Stamens 3 + 3, opposite the perianth-segments, inserted at their base or in the tube; filaments free, partly fused with the perianth or united into a cup (false corona) at the base; anthers basifixed (not in East Africa) or dorsifixed, often versatile, introrse, opening lengthwise by slits

Morphology Reproductive morphology Flowers Gynoecium

Ovary 3-locular, each locule with (1–2–) many axile ovules; style long and slender, with capitate or 3-lobed stigma

Morphology Reproductive morphology Fruits

Fruit a capsule, often loculicidal, or a berry

Morphology Reproductive morphology Seeds

Seeds globose or flattened, sometimes winged, with fleshy endosperm and small embryo

Dutilh, J.H.A. (2009). Neotropical Alliaceae.

Morphology

Description

Perennial geophytes, with a bulb , rhizome or corm , often with alliaceous odour. Leaves concentrated basally, spirally arranged, often forming closed sheaths below, linear , filiform , flat, canaliculated, or terete , with parallel veins and internally with lacticifers. Inflorescence on a terete scape, usually umbel -like, with one to several flowers subtended by 1 (generally) or 2 or more membranous spathe bracts enveloping the immature inflorescence , generally free and spreading, but sometimes united at base, branches and pedicels sometimes also subtended with small bracts. Pedicels not articulate . Flowers hermaphrodite , usually actinomorphic , rarely zygomorphic to resembling an insect. Tepals usually 6, in two whorls apparent or not, rarely less, sometimes 3, with loss of inner ones, petaloid , united at base, rarely free , greenish, white, yellow, rose coloured or purple, rarely ornamented, sometimes with scales at the base of the tepals. Stamens usually 6, in two whorls, sometimes 3 staminodes and 3 fertile, or only 2 fertile. Filaments inserted on the base of the tepals, free or united at the base forming a tube around the ovary . Sometimes with scales or corona between the tepals or filaments.  Ovary superior , tri-carpellate and tri-locular, 1- several ovules per locule , placentation axillary , nectaries septal with openings around the base of the style , or absent. Style at apex of ovary , solitary, erect , solid or sometimes partially hollow near the stigma . Stigma capitate , trilobate, rarely branched. Fruit loculicidal capsule , with 1- several seeds per locule . Seeds flat and dark to black due to phytomelan on the testa.

Distribution

Distribution in the Neotropics

Genera from South America:

• Gilliesia Lindl.: Two species in Chile.
• Ipheion Raf.: Genus with 3 species in Central Chile, Argentina, Uruguay and South Brazil.
• Leucocoryne Lindl. Fifteen species in Chile.
• Miersia Lindl.: Bolivia and Chile.
• Nothoscordum Kunth: This is the biggest genus of the family in South America with about 30 species found in Argentina, Uruguay, Paraguay, Chile, Bolivia, Peru, and Brazil.
• Schickendantziella Speg.: Monotypic genus from Tucumán, Argentina.
• Speea Loes.: Two species in Chile.
• Solaria Phil.: Five species in South Chile and South Argentina.
• Tristagma Poeppig: Genus with about 14 species in Chile and Argentina.
• Trichlora Baker.: Two species in Peru.

Diagnostic

Key differences from similar families

• The Alliaceae can be separated from Amaryllidaceae morphologically mainly by its superiorovary, as opposed to superior in Amaryllidaceae.
• It can be separated from Agapanthaceae by its radially arranged leaves, as opposed to strongly distichous in Agapanthaceae.

Notable genera and distinguishing features

See above.

Useful tips for generic identification

• Gilliesia Lindl.: plants with 1-2  leaves linear, flat. Inflorescence with 2-9 flowers, spathe bracts 2. Flowers bilaterally symmetric, green or purple, sometimes resembling an insect with wings and legs, tepals 6 or rarely 5, free, unequal, with appendages, inner surface with thick epidermis with papillate cells. Stamens 3 fertile, 3 staminodes on a synandrium. Corona of irregular scales on the outside of the staminal cup. Style trilobed.
• Ipheion Raf.: plants sometimes with alliaceous odour. Scape geotropic, decumbent at end of flowering, spathe bracts 2 united for less more than ¾ at one margin and about 1/3 on the other. Flower generally one, actinomorphic, yellow, tepals united at lower 1/3 to ½, filaments free from each other, much longer than the tepal tube, adnate to the perigone in two series, no corona or scales, stigma small capitate. Testa of seeds pitted.
• Leucocoryne Lindl. plants sometimes with alliaceous odour. Scape erect with 1- 12 flowers, spathe bracts linearlanceolate, floweractinomorphic, 6 tepals, isomorphic, united at the base in a conspicuous tube, stamens 3 or 6 fertile, when 3 then opposite the outer tepals, the staminodes long, protruding from the throat. Style short, stigma capitate.
• Miersia Lindl.: inflorescence 3- 7 flowered. Flowers bilaterally symmetric, green with purple lines, with 6 tepals in two series, free, acuminate, with appendages with papillate cells, anthers 6, small, fused in a staminal cup, corona of 6 narrow scales outside, style abaxially reflexed, stigma disk like.
• Nothoscordum Kunth: plants sometimes with alliaceous odour. Scape erect, spathe bracts 2 or rarely more, free or united for less than 1/4. Flowers actinomorphic, mostly white, sometimes with purplish lines, or yellow. Tepals 6 united for less than 1/3, anthers adnate to the tepal tube in one series, no corona or scales, filaments may be united basally, ovary sometimes with small gynophore, stigma small capitate, ovules 1- several per locule, testa of seed smooth. Nothoscordum fictile Macbride and N. sessile Beauverd from NW Argentina differ from all other Alliaceae in South America by having pedicels with individual bracts and by the apiculate outer tepals. These bracts also appear in N. andinum Fuentes, a species sometimes considered as genus Zoellnerallium Crosa.
• Schickendantziella Speg.: inflorescence with 1-2 flowers, flowers actinomorphic, nodding, tepals 3, almost free, caudate, pinkish-brown with purple, anthers 6, filaments connate, enveloping the ovary, no corona or scales. Ovary with many ovules. 
• Speea Loes.: inflorescence 1-3 flowered with 1-2 spathe bracts separate from each other, the outer may form a tube below, flowers pinkish brown with purple, tepals 6, free, caudate, no corona or scales. Anthers basifixed.
• Solaria Phil.: inflorescence short scape with 3-15 flowers, spathe bracts 2. Flowers bilaterally symmetric, green, tepals 6, united. Outer median tepal distinctly larger than the others, giving a labellum like effect, and the inner median tepal often smaller than the two lateral ones. Fertile stamens 1-5, basally connate, staminodes 3 or absent, corona of small scales or absent. Stigma capitate.
• Tristagma Poeppig: plants may have alliaceous odour. Inflorescence scape erect, flowers 1-several, spathe bracts united for less than 1/6, tepals 6 united in a tube for ¼ to 3/5 , anthers 6 included in the tepal tube, filaments absent to long, free from each other, bases partially adnate to the perigone forming small cups, no corona or scales, testa of seeds smooth.
• Trichlora Baker.: bulb tunicated, narrow. Inflorescence with about 5 flowers, spathe bracts 2 free from each other. Flowers actinomorphic, with outer tepals lanceolate, green, free, inner short, obtuse, scale like, green. Stamens 3 fertile, 3 staminodes, corona absent. Stigma with 3 hornlike branches.

General Description

General notes

• The genus Allium L., from the Northern hemisphere i.e. Asia, Europe and North America, has several edible species widely cultivated that may sometimes appear as spontaneous. They can be distinguished mainly by their gynobasic style, i.e. base of style sunken in the middle of the ovary, and by the opening of the nectaries at the base of the ovary.
• Some species from different genera are also cultivated as ornamentals.

Notes on delimitation

• The family Alliaceae, its genera and species have a complex taxonomic history, not yet fully understood.
• The family Alliaceae is closely related to Amaryllidaceae and Agapanthaceae (entirely Old World), with which it is sometimes grouped into one large family. 
• The family is divided in 3 subfamilies: Allioideae, Tulbaghioideae and Gilliesioideae, this last one comprising all South American genera.

Number of genera

• 10 Neotropical genera (see list above).

Status

• Native, and some introduced species.

Literature

Important literature

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

Fay, M.F., Rudall , P.J. & Chase, M.W. 2006. Molecular studies of subfamily Gilliesioideae (Alliaceae). Aliso 22: 367-371.

Guaglianone, R. 1972. Sinopsis de Las especies de Ipheion Raf. y Nothoscordum Kunth (Liliáceaes) de Entre Rios y Regiones Vecinas. Darwiniana 17: 159-240.

Rahn, K. 1998. Alliaceae. In: Kubitzki, K. (ed.) The Families and Genera of Vascular Plants, Monocotyledons - Lilianae (except Orchidaceae) 3:70-78.  Springer-Verlag, Berlin.

Rudall , P.J.,  Bateman, R.M., Fay, M.F. & Eastman, A. 2002. Floral anatomy and systematics of Alliaceae with particular reference to Gilliesia, a presumed insect mimic with strongly zygomorphic flowers. Amer. J. Bot. 81(12): 1867-1883.

Alliaceae, Sarah Smith & Jonathan Stansbie. Flora of Tropical East Africa. 2003

Morphology General Habit

Acaulescent or short-stemmed biennial or perennial geophytes

Morphology Stem

Stem usually swollen to form a bulb or tuberous rhizome, rarely a corm, covered with the dry remains of sheathing leaf-bases

Morphology Leaves

Leaves linear, filiform, lanceolate or rarely ovate, flat, terete or cylindrical and hollow, bases sheathing, veins parallel

Morphology Reproductive morphology Inflorescences Scape

Scape leafless, terete, flat or cylindrical and hollow

Morphology Reproductive morphology Inflorescences

Inflorescence an umbel, subtended by (1–)2 (–several) membranous spathaceous-bracts; bracts enveloping young flower buds; pedicels sometimes further subtended by smaller membranous bracts

Morphology Reproductive morphology Flowers

Flowers hermaphrodite, actinomorphic

Morphology Reproductive morphology Flowers Perianth

Perianth tubular with 6 free lobes; corona present or absent

Morphology Reproductive morphology Flowers Androecium

Stamens usually in 2 series of 3; anthers dehiscing by longitudinal slits

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior, trilocular, with 2 septal nectaries, 2–several ovules per locule; style solitary, erect, at apex of ovary or ± gynobasic; stigma capitate or trilobate

Morphology Reproductive morphology Fruits

Fruit a loculicidal capsule with few-many seeds

Morphology Reproductive morphology Seeds

Seeds small, ovoid, ellipsoid to subglobose or flat