ASPARAGALES often geophytes capsule or berry seed coat obliterated or with

phytomelan
The Asparagales include 14 families, sensu of APG IV 􀇻2016􀇼, although 24 families are
recognized here. This order encompasses a large and diverse number of taxa 􀇻Table 7.2􀇼.
Based on recent phylogenetic studies, it is likely that an apomorphy previously thought to unite
the Asparagales, the presence of seeds having a seed coat containing a black substance called
phytomelan 􀇻Figure 7.29􀇼, may actually be apomorphic for the all except the Orchidaceae,
which is sister to all other members of the order 􀇻Figure 7.30􀇼. The phytomelaniferous seeds of
the Asparagales were lost in some lineages, particularly those that have evolved fleshy fruits.
The phylogenetic relationships of families in the
Asparagales are seen in Figure 7.32. Apomorphies for the
order may include simultaneous microsporogenesis (see
Chapter 11) and an inferior ovary (Figure 7.32); if so, several
reversals in these features occurred in various lineages (Figure
7.32). Family delimitations of the Asparagales have undergone
a number of changes in recent years, and several families
may be united in an alternative classification scheme,

Orchidaceae—Orchid family (orchis, testicle, from the shape

of the root tubers). 700–800 genera/ca. 20,000 species.
(Figures 7.40–7.42)
The Orchidaceae consist of terrestrial or epiphytic, perennial
[rarely annual] herbs [rarely vines]. The roots are often
tuberous (in terrestrial species) or aerial (in epiphytic
species), typically with a multilayered velamen. The stems
are rhizomatous or cormose in terrestrial species, the epiphytic
species often with pseudobulbs. The leaves are spiral,
distichous, or whorled, usually sheathing, simple, and parallel
veined. The inflorescence is a raceme, panicle, spike, or
a solitary flower. The flowers are bisexual, rarely unisexual,
zygomorphic, usually resupinate, resulting in a 180º shift of
floral parts (Figure 7.42C), epigynous. The perianth is biseriate,
homochlamydeous (although outer and inner whorls
are often differentiated), 33, apotepalous or basally syntepalous,
extremely variable in shape and color, sometimes spurred
or with enlarged saclike tepal. The inner median, anterior
tepal (when resupinate; actually posterior early in development)
is termed the “labellum,” which is typically enlarged,
sculptured, or colorful and often functions as a landing platform
for pollinators. The stamen in most species is solitary,
derived from the median stamen of the ancestral outer whorl,
often with two vestigial staminodes derived from the lateral
stamens of an ancestral inner whorl; in Apostasioideae or
Cypripedioideae, there are two or three fertile stamens, when
two, derived from the two lateral stamens of the ancestral
inner whorl, when three, derived from these plus the median
stamen of the outer whorl; the androecium is fused with
the style and stigma to form the gynostemium (also called
the column or gynostegium). Anthers are longitudinally or
modified in dehiscence, bisporangiate, dithecal; in all but the
Apostasioideae and most Cypripedioideae, the pollen is
agglutinated into 1–12 (typically 2 or 4) discrete masses, each
termed a “pollinium” (derived from individual anther
microsporangia or from fusion products or subdivisions of
the microsporangia); the pollinia plus a sticky stalk (derived
from either the anther or stigma) are together termed a
“pollinarium,” the unit of transport during pollination, the
anther connective often modified into an “operculum”
(“anther cap”) that covers the anther(s) prior to pollination.
The pollen consists of tetrad units in most family members,
but may be massulae or monads in various groups (see
Chapter 12, Palynology). The gynoecium is syncarpous, with
an inferior ovary, 3 carpels, and 1–3 locules. The style is solitary
and terminal and is the major component of the gynostemium;
a single, enlarged lobe, termed the “rostellum” and
interpreted as part of the stigma(s), is positioned above the
stigmatic region; the rostellum typically is adherent to the
pollinarium stalk, the tip of which derives a sticky substance
from the surface of the rostellum (this sticky region is termed the “viscidium”). Placentation is parietal or axile;
ovules are
anatropous, usually bitegmic, very many per carpel (sometimes
on the order of a million). Nectaries are typically
present,variable in position and type. The fruit is a loculicidal
capsule or rarely a berry. The seeds are often membranous-
winged, possibly functioning in wind dispersal, and
exalbuminous, the endosperm abortive early in development.
Pollination is effected by various insects (often one species
having a specific association with one orchid species), birds,
bats, or frogs. The transfer of pollen grains together within
the pollinia is an apparent adaptation for ensuring fertilization
of many of the tremendous number of ovules. Some species
have remarkable adaptations for pollination. Among the
more remarkable are several species with visual and chemical
mimicry, fooling a male insect into perceiving the flower as a
potential mate. The bucket orchid, Coryanthes, has an pouchlike
labellum that fills with a fluid secreted from the gynostemium; a bee, falling into this fluid, must travel through a
tunnel, forcing deposition of the pollinarium on its body.
The Orchidaceae were recently classified into five subfamilies:
Apostasioideae (2–3 stamens, axile placentation, lacking
pollinia), Vanilloideae (1 stamen, parietal placentation),
Cypripedioideae (2 stamens, parietal placentation, lacking
pollinia), and the Orchidoideae and Epidendroideae (1
stamen, parietal placentation, pollinia), the last divided by
Cameron et al. (2004, 2006) into a paraphyletic “Lower
Epidendroid” and a monophyletic Higher Epidendroid
(Figure 7.42A,B). The single stamen of the Vanilloideae were
hypothesized by Cameron et al. (2006) to have evolved independently
to that in the Orchidoideae-Epidendroideae (Figure
7.42A,B). Members of the family are distributed worldwide.
Economic importance is largely as cultivated ornamentals,
including some quite monetarily valuable in the horticultural
trade. The fermented capsules of Vanilla planifolia (Figure
7.40B) are the source of vanilla food flavoring. Angraecum
sesquipedale Thouars (Madagascar) is known for its long spur
(up to 45 cm long); this orchid is pollinated by a moth with a
proboscis of that spur length, a fact that Charles Darwin predicted
prior to the discovery of the moth (and recent observation
that it is indeed the pollinator). See Cameron et al. (1999),
Cameron and Chase (2000), and Cameron (2004, 2006) for
recent phylogenetic analyses of the orchids.
The Orchidaceae are distinctive in consisting of mycorrhizal,
mostly perennial, terrestrial or epiphytic herbs having
trimerous, often resupinate flowers with a showy labellum,
the androecium and gynoecium adnate (termed a column,
gynostegium, or gynostemium), the pollen grains often fused
into 1–several masses ( pollinia), bearing a sticky-tipped stalk,
pollinia and stalk termed a pollinarium, which is the unit of
pollen dispersal during pollination.
P (33) A 1–3, when 1 a pollinarium G (3), inferior, with
gynostemium.