®
Floriculture and Ornamental Biotechnology ©2012 Global Science Books
Mexican Geophytes II.
The Genera Hymenocallis, Sprekelia and Zephyranthes
Ernesto Tapia-Campos1 • Jose Manuel Rodriguez-Dominguez1 •
María de los Milagros Revuelta-Arreola1 • Jaap M. Van Tuyl2 • Rodrigo Barba-Gonzalez1*
1 Centro de Investigación y Asistencia en Tecnología y Diseño del Estado de Jalisco A.C. Biotecnología Vegetal, Av. Normalistas #800. Colinas de la Normal., Guadalajara
Jalisco, C.P. 44270, Mexico
2 Plant Breeding, Wageningen University and Research Centre, Droevendaalsesteeg 1, 6708 PB Wageningen, The Netherlands
Corresponding author: * rbarba@ciatej.net.mx; rbarba002@hotmail.com
ABSTRACT
Among hundreds of bulbous ornamental plants native to tropical and subtropical America, different genera from the Amaryllidaceae
family such as Hymenocallis, Sprekelia and Zephyranthes present an enormous potential as ornamental crops. The genus Hymenocallis
comprises over 60 species distributed from the north of Brazil to the south east of the United States; many of them are endemic to Mexico.
The flowers are star shaped and white. The different species grow in a wide and contrasting diversity of habitats, near rivers and streams,
on occasion completely submerged under water and sometimes in dry areas. The genus presents a complicated phylogeny, where in some
cases it is difficult to distinguish species from hybrids. The genus Sprekelia is a monotypic genus native to Mexico; Sprekelia
formosissima is cultivated as an ornamental pot plant in many countries. It presents solitary red flowers, their stems reaches up to 80 to 90
cm. The genus Zephyranthes comprises over 70 species distributed in tropical and subtropical America; different species are cultivated all
over the world as an ornamental crop. The different species have beautiful flowers from white to yellow with various tints from lemon to
sulphur and pink. In this review we will cover taxonomical, chromosomal and phenological aspects of these genera, with the aim of
providing a reference of useful traits for breeding programs.
_____________________________________________________________________________________________________________
Keywords: breeding, in vitro culture, ornamental, perspectives, taxonomy
Abbreviations: BA, 6-benzyladenine; GA3, gibberellic acid; IBA, indole-3-butyric acid; ITS, internal transcribed spacer region; RFLP,
restriction fragment polymorphism
CONTENTS
INTRODUCTION...................................................................................................................................................................................... 129
ORIGIN...................................................................................................................................................................................................... 130
HYMENOCALLIS ...................................................................................................................................................................................... 131
Biology and morphology ....................................................................................................................................................................... 131
Hymenocallis ‘traditional’ classification................................................................................................................................................ 131
Hymenocallis propagation and culture................................................................................................................................................... 131
ZEPHYRANTHES ...................................................................................................................................................................................... 132
Biology and morphology ....................................................................................................................................................................... 132
Zephyranthes propagation and culture................................................................................................................................................... 134
Zephyranthes reproduction .................................................................................................................................................................... 134
Zephyranthes breeding .......................................................................................................................................................................... 135
SPREKELIA ............................................................................................................................................................................................... 136
Biology and morphology ....................................................................................................................................................................... 136
Sprekelia propagation and culture ......................................................................................................................................................... 136
Sprekelia breeding ................................................................................................................................................................................. 136
ALKALOIDS OF HYMENOCALLIS, SPREKELIA AND ZEPHYRANTHES ........................................................................................... 137
CONCLUSIONS........................................................................................................................................................................................ 137
ACKNOWLEDGEMENTS ....................................................................................................................................................................... 138
REFERENCES........................................................................................................................................................................................... 138
_____________________________________________________________________________________________________________
INTRODUCTION
The Amaryllidaceae J. St.-Hil. is a cosmopolitan family
(predominantly pantropical) of petaloid monocots (Meerow
et al. 1999). Its origin is western Gondwanaland in Africa,
from where it radiated to Australia, Eurasia and America
(Huber 1969; Raven and Axelrod 1974; Meerow et al.
1999). The genera Hymenocallis Salisb., Sprekelia Heister
and Zephyranthes Herb. are components of terminal subclades whose higher diversity is present in Mexico. These
Received: 26 January, 2011. Accepted: 20 December, 2012.
genera have been cultivated in some cases, for centuries;
however, it is worthwhile to reconsider their potential as
ornamental plants and for their pharmaceutical importance
due to their alkaloid content. The aim of this work is to give
an insight to the origin of the genera, their culture and
propagation techniques, and to focus attention on traits of
ornamental importance of the Mexican species to set a base
for their use in breeding programs.
Invited Review
Floriculture and Ornamental Biotechnology 6 (Special Issue 1), 129-139 ©2012 Global Science Books
Table 1 Mexican species of the genus Hymenocallis, their distribution in Mexico and some traits of ornamental interest.
Species
Distribution
Traits of interest
H. acutifolia (Herb. ex Sims) Sweet Wide distribution
Narrow foliage, blooms from late summer through autumn
H. araniflora T.M.Howard
Sur Sinaloa, Nayarit
Clusters of long-tubed, sweet-scented blooms, pale green to grayish foliage (Ogden
2007)
H. astrostephana T.M.Howard
Guerrero
Costate leaves, two to five long-tubed flowers (Howard 2001)
H. azteciana Traub.
Jalisco, Nayarit, Zacatecas Spreading, long-stemmed or petiolated leaves with a satiny dusting of gray overlaying
dark green. Small clusters of erect blooms (Ogden 2007)
H. baumlii Ravenna
Chiapas
Erect green star-shaped leaves, ribbon-like flowers (Howard 2001)
H. choretis Hemsl.
C & SW. Mexico
Large-cupped snowy blossoms, with orange pollen. Gray foliage (Ogden 2007)
H. cleo Ravenna
Chiapas
Medium-sized white flowers, broad, glaucous, petiolated leaves (Howard 2001)
H. clivorum Laferr
Sonora
Green stamens, long and thin tepals
H. concinna Baker
Jalisco
Endangered species
H. cordifolia Micheli
Guerrero
Heart-shaped petiolated leaves, umbels of small white flowers (Howard 2001)
H. durangoensis T.M.Howard
Durango
Thick and short sepals. Blooms in early summer. Self-compatible. Endangered specie
H. eucharidifolia Baker
Guerrero, Oaxaca
Broad, green hosta-like foliage. Snowy foliage held in up-facing groups of three or
more
H. galuca (Zucc.) M.Roem.
C&SW Mexico
Large-cupped upright snowy blossoms, with orange pollen with a ammonia or chlorine
fragrance. Gray foliage (Ogden 2007)
H. graminifolia Greenm
Morelos
Dwarf, glaucous leaves, fragrant flowers (Howard 2001)
H. guerreroensis T.M.Howard
Guerrero
Endangered species
H. harrisiana Herb
C & SW Mexico
Small upright flowers with good sized blooms
H. howardii Bauml
SW Mexico
Small plant of glaucous foliage, blooms form mid-June through mid-July
H. imperialis T.M.Howard
San Luis Potosí, Hidalgo Wide, snowy petals. Blooms from late April and May. The bulbs are huge and the big
leaves are spear-shaped.
H. jaliscensis M.E.Jones
Jalisco, Nayarit
Big upright flowers (Borys et al. 2008)
H. leavenworthii (Standl. &
Michoacán
Spreading long stemmed or petiolated leaves with a satiny dusting of gray overlaying
Steyerm.) Bauml
dark green. Small clusters of erect blooms (Ogden 2007). Endangered species
H. lehmilleri T.M.Howard
Guerrero
Elegant white flowers with thin petals, greenish filaments
H. littoralis (Jacq.) Salisb.
Wide distribution
Slender petals attached to the cups of the blooms. Blooms in August. Filaments tipped
with bright orange pollen (Ogden 2007)
H. longibracteata Hochr.
Veracruz
Presumed extinct (Castillo-Campos et al. 2005)
H. maximiliani T.M.Howard
Guerrero
Compact, narrow and erect foliage. Long and narrow sepals
H. partita Ravenna
Chiapas
Aquatic, pleasant fragrance
H. phalangidis Bauml
Nayarit
Daffolil-like plant with few flowered umbels (Howard 2001)
H. pimana Laferr.
Chihuhua
Flowers with yellowish-centered flat cup (Howard 2001)
H. portamonetensis Ravenna
Chiapas
Sharp lanceolate leaves, four to five flowers
H. proterantha Bauml.
SW Mexico
Wide, petiolated glaucous leaves; flowers appear as leaves go dormant (Howard 2001)
H. pumila Bauml.
Jalisco, Colima
Dwarf habit, bright green sword-like leaves (Howard 2001)
H. sonorensis Standl.
Sonora to Nayarit
Clusters of long-tubed, sweet scented blooms, pale green to grayish foliage (Ogden
2007)
H. vasconcelossi Garcia-Mend.
Oaxaca, Puebla
Few oblong-elliptical green leaves. White flowers with thin and short tepals (GarciaMendoza 2010)
H. woelfleana T.M.Howard
Durango, Sonora, Nayarit Presence of a pedicel below the ovary (Howard 1978)
ORIGIN
(Meerow 1987, 1989; Merrow et al. 2002). A study of the
phylogeny of the genus Hymenocallis, based on cladistic
analyses of morphology, plastids and ribosomal DNA,
groups the Hymenocallidae tribe with three genera: Hymenocallis, Ismene Herb. and Leptochiton Sealy, being the
latter sister to the Hymenocallis/Ismene clade. As the genera
Ismene and Leptochiton are endemic to the central Andes
and Hymenocallis is absent from this region, this study
suggests an entry of the Amaryllidaceae through South
America and a vicariance event at some point subsequent to
the origin of the tribe, with a later extinction of the intervening populations of a proto-Hymenocallis ancestor
(Meerow et al. 1999, 2002). Furthermore, Meerow et al.
(1999) investigated the fact that the Eurasian sister clade
has been successful in adapting to temperate habitats, whereas the American clade has not been able to adapt to those
climates. There is also evidence from internal transcribed
spacer regions (ITS) analyses in the genus Zephyranthes of
the probable center of origin of the Amaryllidaceae family’s
extant diversity in the Americas in South America, with a
later migration to North America. The results of Meerow et
al. (2000) show two Zephyranthes clades having originated
by different migration paths northwards from South America. The South American species Z. flavissima Ravenna
and Z. mesochloa Herb. are sister to the Mexican Zephyranthes clade, suggesting a migration from South America
northward through the Panamerican isthmus. The second
clade includes species from Argentina (Z. candida (Lindl.)
Herb., Z. filifolia Baker & Kraenzl.), Brazil (Z. cearensis
Hymenocallis, Sprekelia and Zephyranthes are bulbous
genera of the monocotyledoneous family Amaryllidaceae.
The systematics of the family have undergone study at
several levels (Meerow et al. 1999) and there is a sister
relationship between the Eurasian/Mediterranean clade and
the American genera. The Madrean–Tethyan hypothesis
notes that the initial entry of the family occurred through
North America during the late Cretaceous to the early
Eocene, when North America and western Europe may have
shared a warm, seasonally dry climate (Axelrod 1973,
1975) which might have allowed east/west movement of
species, with island chains of the mid-Atlantic ridge providing stepping stones (Meerow et al. 1999). However,
there are members of the family in Mexico and the United
States that are linked to more basal taxa endemic to South
America (Meerow et al. 1999). The Madrean-Tethyan hypothesis has been questioned after several studies utilizing isozymes, plastid DNA, restriction fragment polymorphisms
(RFLPs), and cladistic analyses of taxa considered emblematic of the disjunction (Little and Chrutchfield 1969;
Kölher and Brückner 1989; Liston et al. 1992; Ray 1994;
Meerow et al. 1999, 2002). Due to the distribution of
Amaryllidaceae in America and the generic richness south
of the equator, a massive extinction event should have
occurred after migration to South America (Meerow et al.
1999). However, there is evidence that the movement of
Amaryllidaceae has been northward from South America
130
Mexican geophytes II. A review of the genera Hymenocallis, Sprekelia and Zephyranthes. Tapia-Campos et al.
Baker), Cuba (Z. rosea (Sprenger) Lindl.) and North America (Z. atamasco (L.) Herb., Z. simpsonii Chapm.), suggesting that this clade entered the southeastern United
States via Cuba after migrating from South America
(Meerow et al. 2000).
Cladistic analyses of plastid DNA sequences rbcL and
trn-L-F and ITS position the genus Hymenocallis as a
monophyletic group; belonging to the Hymenocalliade tribe
along with Ismene and Leptochiton the three genera are enclosed in the Andean clade (Meerow et al. 2000). The genus
Zephyranthes, a polyphyletic group, may represent convergence of two (albeit related) lineages of North (Meso)
American and South America (Meerow 1995), however, a
wider sampling of the genus revealed a putative triple origin
of the genus (Meerow et al. 2000). Zephyranthes, together
with Sprekelia and Habranthus are nested in the Zephyranthinae subtribe of the Hippeastrae tribe, together with the
genera Hippeastrum Herb., Rodophiala C. Presl and Traubia Moldenke, within the Hippeastroid clade.
of this alliance have slightly lanceolate leaves, are usually
evergreen, with shortly joined perigonium segments to the
stamina corona. It is distributed in Colombia, Ecuador and
Mexico. V. Caroliniana, which have introrse anthers, caduceus leaves and a globular ovary with less than four ovules
per locule. These species are native to the southern U.S. VI.
Henryae, distributed in Florida and Cuba, with caduceus
leaves, introrse anthers, oblong ovaries and more than four
ovules per locule.
Hymenocallis propagation and culture
Most of the cultivated Hymenocallis are old heirloom
flowers spread widely by sailors in the 16th century; they
were originally brought from the tropical shores of the
Antilles and the Spanish Main. The identities of these plants
are hidden in a tangled horticultural history. These beautiful
flowers are sold in nurseries as H. littoralis Salisb., H.
caribbaea Herb., H. rotata (Ker Gawler) Herb., or H. tenuiflora Herb. (Ogden 2007). In Mexico, H. laevenworthii
(Standl. & Steyerm.) Bauml, H. proterantha Bauml. and H.
harrasiana Herb. are planted in home gardens and open
areas (Borys et al. 2005a).
Hymenocallis is a very minor component in the bulb
trade, however, the beauty of the spider lily flowers, their
fragrance and their vigorous growth hint at the potential
value of the genus (Hanks and Jones 1987). Some of the
Mexican species are shown in Fig. 1, and a list of traits of
ornamental importance is given in Table 1.
There is little information regarding reproductive capacity of the genus and it is mainly propagated by seeds
produced by self pollination and by bulb offsets (Borys et al.
2005a). Two groups of Hymenocallis can be identified by
their ecology; xerophytes and meso-hydrophytes (Borys et
al. 2005a, 2008). In both groups the seeds lack dormancy;
the xeric group (caduceus leaves) is polyembryogenic, their
seeds show a sequential emergence of embryogenic roots
followed by rapid formation of small rooted, leafless bulbils,
which are forced by contractile roots into deep soil layers
and enter a deep dormancy period. During the following
rainy season the bulbs forms feeder roots and primary
leaves. The meso-hydrophyte group is evergreen and their
seeds produce only one embryogenic root, terminating in a
leafy and rooted bulbil of continuous growth (Borys et al.
2005a).
As mentioned before, traditional propagation of Hymenocallis is done by seeds or by bulb offsets; however,
Leszczynska et al. (2005) utilized scales and rooted basal
plates for the propagation of Mexican spider lilies. Both
scales and the rooted basal plates were able to form bulbils
in a variable number and size per scale or bulb. The bulbs
were produced in different positions, the scales formed
bulbils at the basal end of veins, while in the rooted basal
plates the bulbils were produced either between scales or on
the upper parts of the remaining scales. In the latter case,
the central part had to be removed to avoid the reappearance of the dominant meristem and the bulbils were separated when the basal plate decomposed.
Micropropagation of H. littoralis has been described by
Backhaus et al. (1992). Bulb scales of H. littoralis were
utilized as explants, surface-sterilization was made by submerging the bulbs in a 1% solution of NaOCl, 0.1% Tween®
20 for 20 min, followed by three washes in sterile distilled
water. The scales with a small portion of the basal plate
attached were cultured in darkness in a basal medium (BM)
composed by Murashige and Skoog (1962; MS) salts, supplied with 3% sucrose, 100 mg/l myo-inositol, 0.4 mg/l thiamine-HCl and 7 g/l TC agar; 1-9 bulblets were formed after
5 weeks and were sub-cultured in a proliferation medium
containing the BM medium to which 2 mg/l 6-benzylaminopurine (BA) and 00.02 mg/l indole-3-butyric acid
(IBA) were added. On average, 8 new shoots/bulblet were
produced. After proliferation, the bulbs were placed in a
rooting medium composed of the BM medium plus 6 g/l
agar. The rooted bulbs were successfully transferred to an
HYMENOCALLIS
Biology and morphology
The genus Hymenocallis Salisb. comprises about 70 species
distributed from the southeastern United States to northern
South America, and present in Mexico, Central America
and the Caribbean Islands (Grossi 2007). Mexico possess
the highest number of species of the genus (32, many of
them endemic) (Table 1) with a secondary area of diversity
in the U.S. (Bauml 1979; Smith and Flory 1990, 2001;
Smith et al. 2001).The chromosome number of the genus
has been reported as 2n = 2x = 46 and 40; however, many
aneuploid accessions that have been reported showing a
derivation of these numbers; H. henryae Traub. possesses a
chromosome number of 2n = 2x = 38, being the lowest
reported for the genus (Smith and Flory 1990). Different
species have been cultivated worldwide for two main purposes, their alkaloid content and ornamental value.
‘Spider lily’ is the common name for the genus
Hymenocallis (from Greek “beautiful membrane”); they are
bulbous and perennial herbaceous plants, with elegant,
long-tubed, snowy white flowers, some of them with a
pleasant fragrance. Their bulbs are roundish or oblong, from
which thick whitish roots emerge, covered by a tunic. The
most outstanding trait is the pancratioid morphology of the
flowers, with the stamens united into a distinctive corona
and the long and narrow tepals which give them their characteristic “spidery” look. They have a variable number of
distichous leaves, except H. eucharidifolia Baker, which
has a rosette. The leaves range from 5-7 mm wide and are
sessile, linear and glaucous in H. graminifolia, Greenm. to
15 cm wide, petioled, lanceolate and bright green in H. speciosa Salisb. (Preuss 2002; Grossi 2007).
Spider lilies grow in a wide diversity of habitats. The
North American spider lilies grow in or on the margins of
wetlands habitats, and less often in dry, flat woods or in
disturbed sites, whereas the Mexican spider lilies grow in
most of the climatic conditions of Mexico, from the north to
the south, from xeric to aquatic conditions and from arid
climates with increased elevation to tropical coastal areas
(Preuss 2002).
Hymenocallis ‘traditional’ classification
Traub (1962) grouped the Hymenocallis species into six
alliances based on morphological characteristics. These are:
I. Speciosa, which comprises species with petiolate leaves,
distributed in Brazil, Mexico and the Caribbean. II. Mexicana, with sessile leaves or sub-petiolate that are oblong,
oblong-lanceolate, ensiform or caduceus, native to Mexico.
III. Caribeae, with slightly lanceolate leaves, evergreen,
with free perigonium segments in relation to the staminal
corona. Species of this alliance are distributed in Central
America, the Caribbean and Florida. IV. Littoralis, species
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Floriculture and Ornamental Biotechnology 6 (Special Issue 1), 129-139 ©2012 Global Science Books
Fig. 1 Some Hymenocallis species present in Mexico. (A) H. howardii; (B) H. concinna; (C) H. acutifolia; (D) H. jaliscensis; (E) H. protheranta; (F) H.
laevenworthii; (G) H. concinna in its natural habit. Photos: A. Rodriguez.
intermittent mist greenhouse.
With the application of these techniques it is possible to
achieve massive production for commercial purposes.
Spider lilies are not hardy. H. littoralis prefers a welldrained soil with organic matter and abundant irrigation. In
a study performed by Isdo et al. (2007), 2-year-old bulbs of
H. littoralis were cultivated in a natural field soil (a fineloamy, mixed (calcareous), hyperthermic Anthropic Torrifluvet), in the Sonoran Desert environment of central Arizona, and the plants exposed to two different levels of ambient air of 400 and 700 ppm CO2. This increase in atmospheric CO2 resulted in 48% increase in above ground
growth and 56% increase in bulb growth. Additionally, it
also increased the concentrations of five bulb constituents
that possess anticancer, and antiviral properties.
There have been some attempts to control forcing in H.
x festalis (Worsley) Schmarse (=Ismene x festalis Worsley);
unfortunately, unsuccessfully (De Hertogh and le Nard
1993). Hanks and Jones (1987) stored bulbs of H. x festalis
over winter at 5°C, promoting less flowering. Flower formation in H. speciosa is inhibited in plants held at 25°C but
prompted in plants transferred to 15° or 20°C after 1 July
(Mori et al. 1991). Borys et al. (2009) in an attempt to
break dormancy in H. harrasiana and H. jaliscensis M.E.
Jones stored the bulbs at 18 to 22°C for 5 to 7 months and
planted them in soil at temperatures that ranged from 10 to
35°C. Their results showed that with a longer storage the
time to initiate the emission of leaves decreased, but also
the number of scapes. Anyway, the results varied when the
bulbs were grown in open fields, concluding that neither
clone is suited for forced flowering.
Fig. 2 Some Zephyranthes species present in Mexico. (A) Z. nymphaea;
(B) Z. latissimifolia; (C) Z. fosteri; (D) Z. concolor; (E) Z. macrosiphon;
(F) Z. clintiae. Photos: A. Rodriguez
ZEPHYRANTHES
many of which are endemic. Together with the genus Habranthus they are commonly known as ‘rain lilies’ because
the flowers appear in spring through fall after the first rains
(Knox 2009). The genus takes its name after the Greek god
of the west wind that typically brought the rainfall ‘Zephyrus’. Rain lilies are noteworthy for the wide ecological
Biology and morphology
The genus Zephyranthes Herb. is a bulbous perennial of the
Amaryllidaceae family. It comprises about 90 species, of
which at least 37 are native to Mexico (Table 2; Fig. 2),
132
Mexican geophytes II. A review of the genera Hymenocallis, Sprekelia and Zephyranthes. Tapia-Campos et al.
Table 2 Mexican species of the genus Zephyranthes, their distribution in Mexico and traits of ornamental interest.
Species
Distribution
Traits of interest
Zephyranthes arenicola
Baja California, Isla
Bloom color: white / near white
Brandegee
Guadalupe
Bloom time: late summer / early fall; mid fall.
Zephyranthes bella
Zacatecas, San Luis Potosí
Bloom color: pink; white / near white
T.M.Howard & S.Ogden
Bloom time: late summer / early fall; mid fall.
Zephyranthes brevipes Standl.
Chiapas, Coahuila, Hidalgo,
Bloom color: pink
Morelos
Ovary almost sessile
Zephyranthes carinata Herb.
Mexico to Colombia
Scape bearing a single pink flower
Zephyranthes chichimeca
NE Mexico
Bloom color: pink or rose blushed petals, clustering habit and obovate to spatulate
T.M.Howard & S.Ogden
petals
Bloom time: late spring / early summer.
Zephyranthes chlorosolen
Kansas to E. Mexico, Yucatan Flowers erect; perianth white, sometimes tinged or veined pink
(Herb.) D.Diettr
Bloom time: late spring / early fall (May-Oct).
Zephyranthes chrysantha
Texas to Tamaulipas and
Bloom time: late summer / early fall; mid fall
Greenm. & C.H.Thomps
Nuevo Leon
Z. citrina Baker
SE Mexico, Cuba to haiti
Flowers erect; perianth lemon yellow, funnelform, 3.1–5 cm; perianth tube green.
Bloom time: summer (Jul-Sep)
Zephyranthes clintiae Traub
Querétaro, San Luis Potosí
Bloom color: pink
Zephyranthes concolor (Lindl.) Mexico
Bloom color: chartreuse (yellow-green); white / near white
Benth. & Hook.f
Bloom time: late summer / early fall; mid fall
Zephyranthes conzattii Greenm. Oaxaca
Endangered specie
Zephyranthes crociflora
Coahuila
Bloom color: pale pink; white / near white
T.M.Howard & S.Ogden
Bloom time: late summer / early fall; mid fall
Zephyranthes dichromantha
San Luis Potosí
Bloom color: pale yellow
T.M.Howard
Bloom time: mid summer; late summer / early fall
Zephyranthes drummondii
NW. Florida to NE. Mexico
Flowers erect; perianth white, sometimes flushed pink abaxially, more so with age,
D.Don in R.Sweet
broadly funnelform. Flowering mid spring--mid summer (Mar--Aug).
Zephyranthes erubescens
NE. Mexico
Bloom color: red; white / near white
S.Watson
Bloom time: late summer/early fall; mid fall.
Zephyranthes fosteri Traub
San Luis Potosi and Jalisco to Bloom color: flowers erect or slightly inclined, white, pink or red
Guerrero, Puebla and Oaxaca. Bloom time: late summer / early fall; mid fall.
Zephyranthes howardii Traub
Nuevo Leon, San Luis Potosi
Bloom color: yellow
Zephyranthes katheriniae
Hidalgo
Bloom color: pink; rose / mauve; red; bright yellow
L.B.Spencer
Bloom time: late summer / early fall; mid fall
Zephyranthes latissimifolia
Jalisco
Bloom color: pink; white / near white
L.B.Spencer
Bloom time: late spring / early summer
Zephyranthes leucantha
Hidalgo
Bloom color: white / near white
T.M.Howard
Bloom time: late summer / early fall; mid fall
Zephyranthes lindleyana Herb. NE Mexico to C. America
Bloom color: magenta (pink-purple)
Bloom time: late spring / early summer
Zephyranthes longifolia Hemsl. SE. Arizona to W. Texas and
Bloom color: flowers erect to slightly inclined; pale yellow
Mexico
Bloom time: late summer / early fall; mid fall
Zephyranthes macrosiphon
NE. Mexico
Bloom color: pink; rose / mauve
Baker
Bloom time: mid fall
Zephyranthes minuta (Kunth)
Mexico to Guatemala
Bloom color. pink, rose
D.Dietr.
Big flowers
Veracruz
Bloom color: pink; white / near white
Zephyranthes miradorensis
Bloom time: late summer / early fall; mid fall
(Kraenzl.) Espejo & López-Ferr.
Zephyranthes moctezumae
San Luis Potosi
Bloom color: pale pink especially on the outside of the petals
T.M.Howard
Bloom time: late summer / early fall; mid fall
Zephyranthes morrisclintii
Nuevo Leon
Bloom color: pale pink
Traub & T.M.Howard
Bloom time: late spring / early summer
Zephyranthes nelsonii Greenm. Estado de Mexico, Veracruz,
Bloom color: pale pink, sometimes with red stripes on the petals; white / near white
Oaxaca, Chiapas
Bloom time: late summer / early fall; mid fall
Zephyranthes nervosa Herb.
SE. Mexico
Bloom color: white / near white
Bloom time: late summer / early fall; mid fall
Zephyranthes nymphaea
Tamaulipas, San Luis Potosi
Bloom color: chartreuse (yellow-green)
T.M.Howard & S.Ogden
Bloom time: late summer / early fall; mid fall
Zephyranthes orellanae
Yucatan
Flower erect, actinomorphic. Clear yellow, with or without pale reddish tinges toward
Carnevali, Duno & J. L. Tapia
the apices of the external faces of the external tepals. Species in critical condition.
Zephyranthes primulina
San Luis Potosi
Bloom color: pale or light yellow flowers backed pink-peach, most noticeable when in
T.M.Howard & S.Ogden
bud.
Bloom time: late spring / early summer
Zephyranthes pulchella J.G.Sm. Texas to NE. Mexico
Bloom color: flowers erect, bright yellow with orange or red tints on the petals
Bloom time: late summer / early fall
Zephyranthes reginae
San Luis Potosi
Bloom color: pale yellow, coral / apricot
T.M.Howard & S.Ogden
Bloom time: mid summer.
Rarely seen in cultivation.
Zephyranthes sessilis Herb.
State of Mexico and Distrito
Bloom color: white, white with pink, white with reddish of the external faces of flower
Federal
Zephyranthes subflava
San Luis Potosi
Bloom color: white / near white; cream / tan
L.B.Spencer
Bloom time: mid summer; late summer / early fall
Zephyranthes traubii (W.Hayw.) SE. Texas to NE. Mexico
Bloom color: pale pink; white / near white
Moldenke
Bloom time: late summer / early fall
133
Floriculture and Ornamental Biotechnology 6 (Special Issue 1), 129-139 ©2012 Global Science Books
niche they occupy, from xeric to temporarily flooded conditions, and have many coveted ornamental characteristics
(Knox 2009).
Zephyranthes bulbs are rounded or ovoid, covered with
a thin brown or black tunic. Depending on the species, the
bulbs range in size from 2.5 cm to more than 5 cm in diameter (Knox 2005), the leaves are linear form bright grassy
green to rather broad and glaucous, the size of the leaves
varies from tiny in Z. jonesii (Cory) Traub to broad in Z.
lindleyana Herb.; the scape is slender, hollow and leafless
with one (rarely six) flower(s) at the apex. Rain lily flowers
appear in spring, summer, or fall, depending on the species
(Knox 2005). The flowers of Zephyranthes are funnelshaped and have six petals; they are white, pink or yellow
and crocus-like (Marta 2005). Flowers of novel hybrids
occur in shades of pink, orange and red; some are multicolored with striped or picotee patterns (Knox 2009). Some
species have a sweet pleasant fragrance, which appears to
be recessive; however, there are some species that carry the
trait (Z. drummondi D.Don, Z. morrisclintii Traub &
T.M.Howard, Z. jonesii) (Marta 2005; Roy Chowdhury
2006).
Zephyranthes propagation and culture
The flowers of Zephyranthes lasts just one or two days,
depending on sunlight and temperature, however, new
flowers continuously develop for several days, creating
flushes of flowering. Rainfall triggers blooming (Knox et al.
2009). Most species under cultivation will bloom without
the naturally imposed drought and wet that occurs in nature.
When grown in greenhouses, rain lilies will bloom constantly; however, they have cycles of flowering. Z. primulina
T.M.Howard & S.Ogden might be one of the longest
blooming of all species (Marta 2005). Depending on the
species, a cycle of dryness followed by watering encourages
flowering.
Scagel (2003) planted bulbs of different Zephyranthes
and Habranthus species (Z. candida (Lindl.) Herb., Z. citrina Baker, H. robusta Herb. ex Sweet) into cylindrical pots
containing either a steam-pasteurized or a non-pasteurized
1:1 mixture of Willamette Valley alluvial silt loam and river
sand (11 mg kg-1 available (Bray) phosphorus (P), pH of
6.3).
The pasteurized and non-pasteurized soil was inoculated with the vesicular-arbuscular mycorrhizal fungi
(VAMF) (Glomus intraradices Schenck & Smith), at a rate
of 1:166 (v/v), placing the inoculum at the base of each
bulb at planting; as a control, sterilized inoculum was added
at the base of the bulbs. The plants were maintained in
glasshouses, fertilized and watered as needed. The results of
Scagel (2003) showed that shoots of inoculated plants
emerged 7-13 days before non-inoculated plants; in the case
of Z. candida and H. robusta the emergence of flower bud
was slightly delayed, and flower production was only increased when plants were grown in pasteurized soil; Z.
citrina flowered 4-11 days earlier and the number of
flowers was increased compared to non-inoculated plants;
the leaf biomass was larger for Z. citrina but lower in Z.
candida and H. robusta when the plants were inoculated. In
a second growing cycle, the combined weight of bulbs and
offsets was increased in Z. candida and Z. citrina by 50% to
150% at the end of the growing cycle.
The viability of the seed is lost rapidly; germination
occurs in two to six weeks; under ideal conditions, seedlings will flower in 8 to 12 months (Knox 2009), but 2 years
from seed to flower are reported (Smith et al. 1999). For
this reason, rain lilies are traditionally propagated by bulb
division or seeds; however, this conventional propagation is
slow, seasonal and inconsistent. Additionally, hybrids may
vary in the numbers of offsets they produce (Smith et al.
1999; Knox 2009; Gongopadhyay et al. 2010).
To achieve massive production of bulbs, a viable alternative to conventional propagation is micropropagation.
Disinfestation is a crucial step for in vitro cultures, and on
134
many occasions a considerable amount of plant tissue is
needed because in every disinfestation process, some tissue
must be discarded. The bulbs of some Zephyranthes species
are small, and surface-disinfestation is difficult due to contamination, therefore, different tissue sources should be
utilized. With this aim, Smith et al. (1999) established in
vitro culture from seeds of different Zephyranthes species;
the seeds were surface-sterilized for 2 min. in 95% alcohol,
followed by 20% bleach for 15 min. and rinsed three times
with sterile water. The seeds were germinated in a media
containing MS inorganic salts and 30 g/l sucrose. The bulbs,
twin scales and cross sections of the leaves, as well as the
leaves from the aseptically germinated plants, served as
explants for multiplication and adventitious regeneration.
Bulbs were produced from the twin scale explants and embryogenic callus was obtained.
Aware of the contamination problems for micropropagation in the genus Zephyranthes, Gongopadhyay et al.
(2010) collected bulbs of Z. grandiflora Lindl. (= Z. minuta
D. Dietr.) and washed them thoroughly under running tap
water, removed the outermost scale and immersed them in
Teepol for 2-5 min. with continuous shaking, then rinsed
them in distilled water. Later the bulbs where surface-sterilized with 0.1% HgCl2 for 2 s, washed 3-4 times in sterile
water, cut in segments and placed in MS media supplemented with 3% sucrose and 1 mg/l 6-benzylaminopurine (BA).
The decontamination protocol was unsuccessful and had to
be discarded due to microbial contamination. In a later
attempt to decontaminate the Z. candida bulbs, Gongopadhyay et al. (2010) repeated the decontamination protocol,
nevertheless, this time they treated the bulbs with 0.2 %
Bavistin® and 0.1% Pantomycin for 2, 3, 4, and 5 h under
continuous shaking before surface sterilization with HgCl2.
The antibiotic addition proved to be efficient and they obtained a 67% (4-h treatment) of non-contaminated explants.
The in vitro-established bulbs were utilized for further
micropropagation experiments and two types of explants
were used: i) segmented bulbs and ii) bulb scales from the
inner 2-3 layers. The explants were transferred to MS media
supplemented with 3% sucrose and 2 mg/l BA. From the
basal half of the bulb scales 11 shoots in average were regenerated and more shoots emerged on a basal achlorophyllous bulbous part. The shoots were transferred to MS
media augmented with 2 mg/l BA and 1 mg/l gibberellic
acid (GA3) for enhancement. Stout roots were developed in
the presence of 1 mg/l indole-3-butyric acid (IBA) and the
plants were successfully transferred to the greenhouse.
Zephyranthes reproduction
Different chromosome numbers have been reported for different species of Zephyranthes, which varies even among
the same species (Table 3). The different chromosome
numbers range from 2n = 18 to 96 (Raina and Khoshoo
1971b) and shows an interesting aneuploid-polyploid
chromosomal polymorphism. Not only different chromosome numbers but even different cytotypes have been
reported for different Zephyranthes species in a detailed
study of chromosome sets accomplished by Coe (1954). He
found different chromosome numbers and cytotypes among
plants of Z. chlorosolen D. Dietr.; in a couple of plants he
found 58 and 59 chromosomes, while in others the chromosome number was 48; interestingly, the chromosomes from
the plants with 58 and 59 chromosomes were smaller than
those with 48 chromosomes and the total length of the
chromosomes in a cell of the 59-chromosome plant was not
greater than those in a cell of a 48-chromosome plant. Furthermore, the chromosome set in a 48-chromosome plant
was not represented by 24 pairs of homologous chromosomes but by 23 homologous pairs and 2 chromosomes
without morphological homologues. Similarly, Z. drummondii D. Don presented chromosome numbers of 24 and
48 in the same plant; different chromosome sets of 48chromosome plants were constituted by 23 and 22 homologous chromosomes and 2 and 4 chromosomes without mor-
Mexican geophytes II. A review of the genera Hymenocallis, Sprekelia and Zephyranthes. Tapia-Campos et al.
Table 3 Chromosome numbers (2n) reported for Zephyranthes species.
Species
Synonym
2n Chromosome numbers
Z. atamasco
24
Z. candida
19, 32, 38, 40
38
38, 40, 41
Z. chlosolen
Z. brazosensis
48, 55, 56, 58, 59
Z. citrina
Z. sulphurea
48
Z. drummondii
Z. pedunculata
24, 48
Z. longifolia
24, 44, 45, 46, 48, 49, 50
Z. aff. mesochloa
12
Z. minuta
Z. grandiflora
24
24
24
48
Z. puertoricensis
Z. tubispatha
24
25
Z. rosea
24
Z. aff stellaris
20
Z. taubii
24
phological homologues, respectively.
The most probable basic number of the genus is x = 6.
(Flory 1968; Raina and Khoshoo 1971; Flory 1977;
Meerow et al. 1999) although x = 5 has also been proposed
(Daviña and Fernandez 1989). To determine whether 6 or 5
is the basic chromosome number of the genus, meiotic studies had to be performed, though meiotic analyses are
scarce due to the difficulty of obtaining meiotic cells as the
flowers develop inside the bulbs and the bulb has to be
destroyed to obtain the flower buds and it is not possible to
know in advance the meiotic stage of the cells.
In a cytological study of reproduction in Z. drummondii
(Coe 1953) the microsporogenesis analyses revealed several
abnormalities such as precocious chromosomes, bridges and
fragments, non-disjunctions and chromosome eliminations;
despite all the irregularities observed, the pollen fertility
was about 91%. The more interesting abnormality was the
clumpling of multivalent formations, containing more than
20 chromosomes in some cases. Tohibi Devi and Borua
(1997) reported trivalent, quadrivalent and pentavalent
associations in microsporogenesis analyses of Z. minuta D.
Dietr. and Z. candida and low fertility in pollen germination.
It seems that such abnormalities are common in Zephyranthes species; in a study of the pollen grain mitosis in Z.
puertoricensis, Traub the chromosome numbers ranged
from n = 1–16 (Raina and Khoshoo 1971), where those with
less than 6 chromosomes might not be fertile, but those with
6 or more might be able to generate odd chromosome numbers. In the genus Lilium (2n = 2x = 24), triploid hybrids
(2n = 3x = 36) were able to produce progeny, however, the
chromosome numbers of the progeny was, in some cases,
aneuploid, due to the production of pollen with odd chromosome numbers (Barba-Gonzalez et al. 2006). Despite all the
abnormalities in meiosis, the pollen fertility does not seem
to be affected. In a pollen fertility screening of different
cytotypes of Zephyranthes whose chromosome numbers
ranged from 24 to 73, all of them were fertile to some
degree, from a maximum of 95% in those cytotypes with 48
chromosomes to 30% in those with 24 (Raina and Khoshoo
1972a). Kapoor and Tandon (1963) analyzed the endosperm
formation in Z .minuta (2n = 24) (= Z. grandiflora; World
Check List of Selected Plant Families) which had an
endosperm of 3n = 36.
Cytological studies performed in Z. drummondii D.Don
(= Cooperia pedunculata Herb.); World Check List of
Selected Species) showed that it is a pseudogamous apomictic and that the occurrence of semigamy is frequent.
Pollination is a pre-requisite for the setting of seed (Coe
1953).
In addition to the apomictic condition of some Zephyranthes it seem that species with styles longer than the
filaments are self-incompatible, those with styles shorter
than the filaments are self-compatible and those with styles
135
Reference
Radford et al. 1968
Tohibi Devi and Borua 1997
Yokouchi 1965
Raina and Khoshoo 1972a
Coe 1954
Raina and Khoshoo 1972c
Coe 1954
Coe 1954
Daviña and Fernandez 1989
Tohibi Devi and Borua 1997
Kaboor and Tandom 1963
Raina and Khoshoo 1972a
Coe 1954
Raina and Khoshoo 1972a
Raina and Khoshoo 1971
Raina and Khoshoo 1972a
Daviña and Fernandez 1989
Coe 1954
more or less equal to the length of the filaments are selfincompatible (Raina and Khoshoo 1972c). However, this
seems not to be the rule as in the case of Z. atamasco, a
long-styled species is reported to be fully self-compatible
(Broyles and Wyatt 1991).
One of the closest relatives of Zephyranthes is the genus
Habranthus (Meerow et al. 1999) where chromosomal
numbers of 12 and 18 are found (Felix et al. 2008). Considering these numbers, the aneusomaty (Coe 1954), the
multivalent formation and all the abnormalities during
meiosis (Coe 1953; Tohibi Devi and Borua 1997) present in
different species of the genus Zephyranthes, it is clear that
the genus has polyploidy, ranging from 2x to 16x with a
basic number of x = 6 (Raina and Khoshoo 1972a). All this,
coupled with self-compatibility / incompatibility and sexual
apomictic reproduction, together with hybridization and
polyploidy, the production of sex cells with a wide range of
numbers, tolerated due to the inherent polyploidy constitution, has lead to a cycle of aneuploid variability, with a
genetic system in the genus with the potential to conserve
and preserve all heterozygosity through agamospermy and
in particular through vegetative multiplication (Raina and
Khoshoo 1972c).
Zephyranthes breeding
There are many traits that are pursued in Zephyranthes
breeding, these include ruffled petals, bronze foliage, fragrance, colors of varying intensity (ranging from apricot to
salmon, orange, cherry red and purple), stripes, picotees,
bicolours as well as flowers that remain open for as long as
3 days (RoyChowdhury and Hubstenberger 2006)
Many Zephyranthes species are apomictic [Z. primulina,
katheriniae L.B. Spencer, macrosiphon Baker, citrina, longifolia Hemsl., pulchella J.G. Sm. are apomictic (Roy
Chowdhury and Hubstenberger 2006)]. This trait is inheritable (RoyChowdhury and Hubstenberger 2006), and is seen
as a desirable character in breeding programs, because once
a cultivar is established it will produce many seeds and the
progeny will be genetically identical to the mother plant.
On the other hand, when breeding for new traits, the apomictic species have a drawback when used as female
parents because the crosses will always be successful, but
never produce hybrid seed (RoyChowdhury and Hubstenberger 2006). For this reason, identified apomictic species
are utilized as male parents.
There are many Zephyranthes intra- and interspecific
hybrids. Perhaps the most well known hybrid is ‘Ajax’,
which is a cross between Z. candida x Z. citrina (Knox
2009), but is not an ‘easy to find’ hybrid these days. After
‘Ajax’ another popular hybrid is the trihybrid ‘Grandjax’
from the cross between Z. grandiflora x Z. ‘Ajax’. Like
these hybrids, many others have been produced, but in most
Floriculture and Ornamental Biotechnology 6 (Special Issue 1), 129-139 ©2012 Global Science Books
of them the lineages are not clear. There are a few hybridization programs that can be mentioned, among them are
the hybrids developed by RoyChowdhury and Hubstenberger (2006), who utilized 44 different female parents, including botanical species and hybrid (some of them interspecific) and trihybrid cultivars such as [(Z. candida x Z.
citrina) x Z. macrosiphon). The 44 female parents were
crossed with 46 male parents, in total 215 crosses were
performed of which 41% (87) were successful. Within these
crosses 34 out of the 44 female parents made at least one
fertile cross. Chu (2009) produced a series of interesting
hybrids, where different color combinations were obtained
in which selfing of different species was proven in the production of these hybrids; Z. citrina yielded seeds in 100%
of the self-pollinations, while Z. rosea, Z. candida and Z.
grandiflora (= Z. minuta; World Checklist of Selected
Families) produced seeds in 22.6, 5.3 and 1.1% of the
crosses, respectively. Successful interspecific crosses included Z. candida x Z. citrina; Z. citrina x Z. candida, Z.
minuta and Z. rosea. Selfing of the F1 generation were
performed with success. BC1 crosses were performed as
follows: Z. candida x (Z. candida x Z. citrina); (Z. citrina x
Z. minuta) x Z. candida and (Z. citrina x Z. minuta) x Z.
citrina. However, it is noticeable that in most of the crosses
involving Z. citrina as maternal parent, the progeny resembled the maternal phenotype, suggesting that these plants
might not true hybrids, but a product of the apomictic
nature of Z. citrina.
Fig. 3 Sprekelia formossisima. Photo: A. Rodriguez.
SPREKELIA
Biology and morphology
Sprekelia Heist. (2n = 60, 120) is an herbaceous monocotyledonous genus of perennial bulbs of the Amaryllidaceae
commonly known as ‘Aztec lily’ and ‘Jacobean lily’. The
genus comprises two recognized species (World Checklist
of Selected Species 2011) viz. Sprekelia formosissima (L.)
Herb. and Sprekelia howardii Lehmiller; nevertheless other
species are mentioned in literature, however it may be a
monotypic genus and the mentioned species are variants of
S. formosissima (Fig. 3) (Rodriguez-Contreras A, Instituto
de Botanica y Zoologia, Universidad de Guadalajara,
Mexico, pers. comm.). It is cultured worldwide as a pot
plant. Its introduction to Europe can be traced back to the
end of the 16th century, when it was known as ‘Narcissus
Indicus Jacobeus’ (Ramon-Laca Menendez de Luarca 1997).
It has a zygomorphic perianth adapted for hummingbird
pollination (Meerow et al. 2000). The flower stems are 4070 cm high, sometimes reaching 90 cm; they are orchidshaped and occur in various shades of red (Howard 2001),
often with a white line in the center of the tepals and sometimes with white borders on tepals 1, 3 and 5. Two groups
have been described, the first with wide perennial leaves
and the second with thin deciduous leaves (Borys et al.
2005b). The leaves are sometimes glaucous.
It is distributed from the North of Mexico from Chihuahua to the south in the state of Oaxaca (Howard 2001).
It belongs to the Zephyranthinae subtribe of the Hippeastrae tribe, of the American clade. It is a sister genus to
Zephyranthes to which it is closely related (Meerow et al.
1999).
Sprekelia propagation and culture
Plants of Sprekelia grow in a wide variety of soils, from
deep to shallow, clayish, sandy or rocky, and from temperate to dry and hot climates (Borys et al. 2005b). Naturally it grows among xeric bushes, among rocks in oak
forests and disturbed places, from 750 to 2700 m (OrtizMedina 1996). Flowering of Sprekelia is reported to occur
from April to July, however, under irrigation it can be extended to August, (Borys and Leszczynska-Borys 1999).
The bulbs have a dormancy period in winter and spring,
which can be overcome by dry storage for a few weeks after
136
flowering (Lozano 1992).
The traditional propagation of Sprekelia is through bulb
division and occasionally by seeds; bulb division is
preferred because bulb maturity takes up to four years when
grown from seed. An average of 85% germination of S.
formosissima seeds was reported in temperatures ranging
from 10-35°C, it took four days at temperatures ranging
from 20-25°C, however, at extreme temperatures (10-15°C
and 30-35°C) the germination percentage decreased slightly
and the days to germinate increased (Borys et al. 2005c).
Besides traditional propagation, micropropagation of
Sprekelia has been achieved with 83% survival utilizing
small bulbs established in vitro, cultured in media containing MS inorganic salts, supplemented with 100 mg L-1
myo-inositol, 0.4 mg L-1 thiamine, 1.0 mg L-1 pyridoxine,
1.0 mg L-1 niacin, 0.4 mg L-1 glycine, 80 mg L-1 adenine
sulphate, 20 μm 6-benzyladenine (BA) and 3% sucrose.
The bulblet growth was promoted by subculturing the
bulblets in the same media without the adenine sulphate and
the BA, with a sucrose concentration of 5%. The bulbs were
rooted in a medium containing MS inorganic salts, supplemented with 100 mg L-1 myo-inositol, 0.4 mg L-1 thiamine, 1.0 mg L-1 pyridoxine, 1.0 mg L-1 niacin, 0.4 mg L-1
glycine, 0.2 mg L-1 IBA and 3% sucrose (Cazarez-Prado et
al. 2010).
Sprekelia breeding
Several Sprekelia cultivars have been released to the market,
‘Orient Red’ is considered to be one of the best cultivars,
which flowers with the first rains and is able to flower again
in the fall (Howard 2001). Besides the intra-specific hybrids,
perhaps one of the most interesting features of Sprekelia is
the relative ease that it is hybridized with other genera.
Several intergeneric hybrids have been reported, such as i)
Hippeastrum x Sprekelia = x Hippeastrelia, whose chromosome number has been doubled in vitro to restore fertility
(Van Tuyl pers. comm.); among this group there are complicated crosses such as (H. papilio (Ravenna) Van Scheepen x H. fragrantissimum (Cárdenas) Meerow) x Sprekelia
formosissima (Fellers 1998); ‘Mystique’ is a remarkable
hybrid because the plant sets fertile seeds (Howard 2001)
and Sprekelia x Habranthus = x Sprekanthus Traub.
Recently, another intergeneric hybrid has been reported as x
Sprekelianthes Lehmiller, this is the result of a cross of
Sprekelia formosissima x Zephyranthes traubii; the bigene-
Mexican geophytes II. A review of the genera Hymenocallis, Sprekelia and Zephyranthes. Tapia-Campos et al.
(+)-8-O-demethylmaritidine
1-O-acetylpseudolycorine
4,5-dehydro-anhydrolycorine
4,5-dehydro-anhydropseudolycorine
7-deoxynarciclasine
7-deoxy-trans-dihydronarciclasine
11-Hydroxyvittatine
N-demethylgalanthamine
Anhydrolycorine
Anhydro-pseudolycorine
Caranine
Galanthamine isomer
Galanthine
Haemanthidine
Haemanthamine
Hippeasterine
Lycoramine
Lycorine
Narciclasine
Narcissidine
Nerinine
Pancratistatin
Pseudolycorine
Tazettine
H. venezuelensis
H. tubiflora
H. speciosa
H. sonorensis
H. rotata
H. lobata
Hymenocallis species
H. latifolia
H. littoralis
H. harrisiana
H. bolivariana
Table 4 Alkaloids present in different species of the genus Hymenocallis.
Alkaloids
+9
+8
+8
+8
+5,7
+5,7
+9
+8
+8
+5
+8
+8
+8
+8
+8
+8
+2
+
8
+8
+2
+2
+2
+8
+4
+
9
+
4
+8
+1
+1
+1
+1
1
1
+8
+
2
9
+
+5,7
+
+
+6
+6
+
8
+8
+8
+2
+
3,5,7
+2
+2
+
6
+
6
+1
+6
+8
+
4
+8
+
1
+
+8
1
The superscript refers to the bibliographic source as follows: 1 Boit 1958; 2 Boit 1960; 3 Backhaus et al. 1992; 4 Antoun 1993; 5 Petit et al. 1993; 6 Petiti et al. 1995; 7 Idso
et al. 2000; 8 Rivero et al. 2004; 9 Abou-Donia 2008.
ric hybrid resembles Sprekelia, it displays the same characteristics, but its dimension is about half the size of Sprekelia
(Lehmiller 2003-2004). It is argued that the relative ease of
Sprekelia hybridization with the above-mentioned genera is
due to its taxonomic proximity. Meerow et al. (1999)
groups the genus Sprekelia in the Hippeastrae tribe, together
with Habranthus, Zephyranthes, Hippeastrum, Rhodophiala
and Traubia.
ALKALOIDS OF HYMENOCALLIS, SPREKELIA
AND ZEPHYRANTHES
Besides the ornamental importance of the genera, several
genera of the Amaryllidaceae are of pharmaceutical importance due to their alkaloid content. Alkaloids are groups of
complex heterocyclic nitrogen compounds, which have
strong physiological activity, are often toxic, and retain their
own basic chemical properties’ (Aniszewski 2007).
The alkaloids of the Amaryllidaceae are a group of isoquinoline alkaloids, produced almost solely by members of
this family. These alkaloids have attracted considerable
amount of interest due to some important pharmacological
activities they have been shown to possess (Unver 2007).
There are many different L-tyrosine-derived alkaloids
present in the genus Hymenocallis where at least 24 alkaloids have been reported (Table 4) (Boit and Döpke 1958,
1960; Backhaus 1992; Antoun et al. 1993; Pettit et al. 1993,
1995; Idso et al. 2000; Rivero et al. 2004; Abou-Donia et al.
2008); in the genus Zephyranthes there are 18 reported
alkaloids and 7 for Sprekelia formosissima (Table 5) (Pettit
1984; Pettit et al. 1990; Gabrielsen et al. 1992; Hohmann
2002; Mutsuga 2002; Kaur et al. 2007; Unver 2007; Wu et
al. 2009a, 2009b, 2010).
These alkaloids have a wide range of biological activities, they are known to have analgesic, cytotoxic activity
against various human tumoral cell lines, and anti-malarial,
antineoplastic and display effects on the central nervous
system (Aniszewski 2007). Among the different alkaloids
137
present in the genera Hymenocallis, Zephyranthes and Sprekelia the lycorine-type alkaloids are particularly important
for Alzheimer’s disease. Other alkaloids present in the different genera have been useful in the treatment of tumors,
breast cancer, diabetes mellitus and human epidermoid carcinoma as well as other anti-cancer activities (Pettit 1984;
Kaur et al. 2007). Alkaloids from Z. carinata Herb. induced
apoptosis in tumor cells by blocking the progression of the
cell cycle at G0/G1 and S phases; and Z. candida was found
to contain trans-dihydronarciclasine, a cytostatic constituent utilized for leukemia treatment (Pettit et al. 1990).
Also, the alkaloids in these genera present antiviral activity and in vitro inhibition of poxvirus (Gabrielsen et al.
1992; Mutsuga 2002).
In addition to the alkaloids and other substances obtained from bulbs and seeds of the genus Zephyranthes, six
ceramides have recently been identified in Z. candida: candidamide A and B, zephyrantnamide A, B, C and D. These
are the first reported for the Amaryllidaceae. The importance of ceramides lies in its role as signalling molecules
involved, among other functions, in programmed cell death
and apoptosis. Candidamide A and B displayed moderate
activities against bacteria Staphylococcus aureus and Escherichia coli, and fungi Aspergillus niger, Candida albicans
and Trichophyton rubrum (Wu et al. 2009a, 2009b, 2010).
CONCLUSIONS
The genera Hymenocallis, Sprekelia and Zephyranthes offer
great potential as ornamental plants and many cultivars and
hybrids are already present in the market. However, the
addition of Mexican species for breeding programmes
would provide genetic variability resulting in novel cultivars with interesting ornamental traits of importance. These
three genera are related and interspecific hybridization as
well as intergeneric hybridization is possible. The inclusion
of other related genera, such as Habranthus could speed up
the breeding process. The development of micropropagation
Floriculture and Ornamental Biotechnology 6 (Special Issue 1), 129-139 ©2012 Global Science Books
Table 5 Alkaloids present in different Zephyranthes and Sprekelia species.
Alkaloids
Z. candida Z. carinata Z. citrina
l-O-(3-hydroxybutyryl)pancratistatin
l-O-(3-O--O-glucopyranosylbutyryl)pancratistatin
3-epimacrinine
7-deoxynarciclasine
8-O-demethylmaritidine
Crinamine
Crinidine
Epimaritidine
Galanthine
Haemanthamine
Haemanthidine
Ismine
Lycorenine
Lycorine
Maritidine
Narcissidine
Nerinine
Oxomaritidine
Pancracine
Pancratistatin
Pretazettine
Tazettine
trans-dihydronarciclasine
Vittatine
Species
Z. minuta Z. rosea
‘Ajax’ (Z. candida
x Z. Citrina)
Sprekelia
formossisima
+5
+5
+6
+4
+6
+2
+1
+2
+8
+3,8
+3
+8
+7
+7
+2
+7
+7
+7
+2
+1
+6
+6
+6
+1
+1
+1
+3
+7
+7
+8
+4
+3
+4
+1
+6
+6
+7
The superscript refers to the bibliographic source as follows: 1 Boit and Döpke 1961; 2 Ghosal and Razdan 1985; 3 Petit et al 1990; 4 Gabrielsen et al 1992; 5 Kojima et al
1998; 6 Hohmann et al . 2002; 7 Aniszewski 2007; 8 Wu et al 2009
techniques in the different genera should speed the development of new varieties. Basic, as well as applied research
must be performed in order to understand the complicated
chromosomal segregation, especially in the genus Zephyranthes. This research might help to unravel the mystery of
the family origin in the Americas. There are many species
of the different genera that must be domesticated to become
part of breeding programmes; research has been conducted
regarding the phenology of several species, however future
research must be conducted to determine the phenology of
species that are not commercially known.
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ACKNOWLEDGEMENTS
The authors would like to express their gratitude to Dr. Aaron
Rodriguez-Contreras, of the Instituto de Botanica y Zoologia, Universidad de Guadalajara, Mexico for pictures for this work.
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