keys to the vascular plants of east texas - Botanical Research ...

444 AMARYLLIDACEAE
cymes—Müller-Doblies 1977; Meerow & Snijman 1998), sometimes reduced to a single flower,
subtended by bracts; flowers often showy; perianth of 6 similar, usually partly connate, petallike
segments (= tepals), an additional whorl of tepal-like tissue, the crown or corona, sometimes
present within the tepals; stamens 6; filaments separate or connate; ovary inferior, of 3
carpels; style 1; placentation axile; fruit usually a capsule.
AA medium-sized family (59 genera and ca. 850 species) of mostly bulbose, scapose perennial
herbs (Meerow & Snijman 1998), often with showy flowers. While primarily tropical, the species
are widely distributed, with centers of diversity in South America (28 genera, especially
the Andes), Africa (19 genera), and the Mediterranean (8 genera); only Crinum (with water dispersed
seeds) is found in both the Old and New worlds. Such a distribution supports a
Gondwanan origin for the family, dating from a time when the southern continents were in
close proximity (Raven & Axelrod 1974; Meerow & Snijman 1998). Ito et al. (1999), based on
molecular evidence, found that the earliest branch of the family is African. They therefore suggested
that the Amaryllidaceae originated in Africa, with South America as a secondary center
of diversification. The family is rich in distinctive alkaloids, with at least one, pancratistatin,
showing promise as an anticancer drug (Pettit et al. 1993, 1995). The Amaryllidaceae is economically
most important for its ornamentals—Narcissus, Leucojum, and Galanthus are reported
to be “among the most important temperate-zone spring-flowering bulbs in commerce”
(Meerow & Snijman 1998), and Amaryllis, Crinum, Cyrtanthus (KAFFIR-LILY), Eucharis (AMA-
ZON-LILY), Haemanthus (BLOOD-LILY), Hippeastrum, Hymenocallis, Nerine (GUERNSEY-LILY), and
Zephyranthes are among the many other ornamentals (Judd et al. 1999). The genera have been
variously treated in terms of family affiliation. Many authorities have put them in a very
broadly defined and clearly polyphyletic (but practical) Liliaceae (e.g., Cronquist 1988; Diggs et
al. 1999) based on superficial similarities of the flower structure to that of the genus Lilium,
while others have treated them as a separate Amaryllidaceae variously delimited based on particular
characters such as an inferior ovary (e.g., Correll & Johnston 1970) or an umbellate inflorescence
subtended by bracts (e.g., Hutchinson 1934; Traub 1957a). Based on phylogenetic
analyses (e.g., Fay & Chase 1996; Meerow et al. 1999 [2000], 2000b; Chase et al. 2000; Fay et al.
2000), we are following many recent authors (e.g., Mabberley 1997; Meerow & Snijman 1998) in
recognizing the Amaryllidaceae as distinct and excluding such groups as the Alliaceae,
Hemerocallidaceae, or portions of the Agavaceae, which have sometimes been included based
on such characters as inflorescence similarities (e.g., Traub 1963a) or ovary position (Correll &
Johnston 1970) (see further discussion under Alliaceae). As recognized here in a fairly narrowly
defined sense, the Amaryllidaceae is a well-defined monophyletic group in the order
Asparagales and most closely related to the Agapanthaceae, Alliaceae, and Hyacinthaceae (Fay
& Chase 1996; Ito et al. 1999; Meerow et al. 1999, 2000b). As a member of this order, the
Amaryllidaceae is more closely related to families such as the Agavaceae, Iridaceae, and
Orchidaceae than it is to other taxa often put in a broadly defined Liliaceae (Chase et al. 1995a,
1995b, 1996, 2000; Fay et al. 2000). Unfortunately, family circumscription of the
Amaryllidaceae is not yet settled—recently, the Angiosperm Phylogeny Group (APG II 2003)
suggested either submerging the Amaryllidaceae into a very broadly defined Alliaceae or alternatively
recognizing them as a separate family. It is now clear that the Amaryllidaceae and
other members of order Asparagales have evolved many characteristics in parallel with families
in order Liliales (Dahlgren et al. 1985). For a detailed discussion of the groups formerly
treated as Liliaceae in the broad sense, see the family synopsis of the Liliaceae (here treated in a
restricted sense) on page 726. Family name from Amaryllis, BELLADONNA-LILY, a genus now
considered to be represented by a single South African species. (Greek: Amaryllis, a shepherdess,
possibly from a root word meaning sparkling, in reference to the attractiveness of the flowers—
Shosteck 1974) (subclass Liliidae—Cronquist; order Asparagales—APG II)
FAMILY RECOGNITION IN THE FIELD: scapose bulbose herbs with inferior ovaries, 6 stamens, usually
showy flowers, and umbellate inflorescences subtended by bracts.