Introduction Morphological Concepts

In this book we present an integrated account of The following short explanations of terms refer
Les monopétales régulieres constituent moins une famille qu'une to the monocotyledons only and are not meant
the monocotyledons. We have attempted to decide
grande nation dans laquelle on compte plusieurs familles bien distinctes ; whether the character states of the monocotyle- to be general definitions. They apply primarily to
en sorte que pour les comprendre toutes sous une indication commune, dons as a whole, and those of their constituent the concepts used in the chapters on character
il faut employer des caractêres si généraux et si vagues que c'est paraitre groups, are primitive or derived. On the basis of states and their distributions and in the taxonomic
these considerations, and with the use of some jus- section of the book.
dire quelque chose en ne disant en effet presque rien du tout. Il vaut tifiable general assumptions, we present some evo-
mieux se renfermer dans des bornes plus étroites, mais qu'on puisse lutionary models in accordance with DAHLGREN
assigner avec plus de précision. and RASMUSSEN (1983). In doing this we use the Underground Parts
elementary terms currently employed by the school
J.-J. ROUSSEAU of cladistics. Many of the character states and their In monocotyledons the first root formed on the
Lettres sur la botanique. Lettre IV distributions are presented in greater detail by embryo, the radicle, is ephemeral and sometimes
19e Juin 1772. DAHLGREN and CLIFFORD (1982). hardly distinguishable. The root system arises
The major part of the book is taken up by our from the basal nodes of erect shoots or from any
classification of the monocotyledons, which is syn- node in prostrate shoots. Roots produced by aerial
The regular monopetals constitute less a family than a great nation in thetic in the sense that it uses data of many differ- shoots may be green and assimilatory (as are the
which one may recognize several quite distinct families; so that in ent kinds and evolutionary in the sense that the roots of epiphytic orchids and aroids) or may form
order to describe them under a common heading, it is necessary to employ evolutionary model is given as much weight as massive props, as in Pandanus and some palms.
characters so general and so vague that the heading when it appears to say possible. The classification is sufficiently practical, Some of the cells of the root epidermis send out
something is saying in effect almost nothing at all. It would be better we believe, for use in the herbarium and in the root hairs. These root-hair epidermal cells may re-
to restrict oneself within narrower boundaries that can be delimited with field, though for the latter assistance from artificial semble other epidermal cells of the root, but in
greater precision. diagnostic keys will be required as well. Our orders some groups they are conspicuously shorter and
and families are generally rather narrow and the are called root-hair short cells (Fig. 27).
families, consequently, are numerous, especially in The roots in some groups of monocotyledons are
Asparagales. Keys to the families of each order fusiform or tuber-like; these contain nutrients and
have therefore been provided. function as storage roots.
We believe that a concept of Amaryllidaceae, for The underground stem, when well-developed, may
example, which includes Amaryllidaceae sensu be a rhizome, corm or tuber. It is often elongate,
stricto, Ixioliriaceae, Agavaceae, pro parte, and, and either horizontal or vertical, and then forms
perhaps, Alstroemeriaceae, is of no help to the a rhizome; plants with a rhizome are described
botanist seeking to recognise natural and compre- as rhizomatous. A short, compact underground
hensible groups, and that it will prevent him from stem filled with nutrients is a corm, which is
understanding the evolutionary pathways within synonymous with rhizomatous tuber. The corm
and around the family. Again, it is certainly of axis generally extends over several internodes. It
no advantage to unite Liliaceae, as circumscribed may be enclosed by a tunic of dry leaves or leaf
here, with Asphodelaceae, Hypoxidaceae, Teco- bases which sometimes form a characteristically
philaeaceae, and Trilliaceae, if at the same time sculptured fibrous envelope; such tubers are called
the Liliaceae are kept distinct from Orchidaceae, tunicated corms (Fig. 117B), and are particularly
Alstroemeriaceae or Iridaceae. common in Iridaceae. Part of the underground
It has been our intention to avoid the constraints stem may become inflated to form a globose stor-
of convention and to adopt an unbiassed ap- age organ and is then called a fuber. Frequently,
proach, using as wide a range of data as we could as in many Dioscoreales and Arales, the tuber
within the limitations of time (self-imposed), re- seems to be chiefly made up of the axis immediate-
sources and competence. ly below the cotyledon, the Aypocotyl, and is then
termed a hypocotylar tuber.
2 Morphological Concepts Leaves 3

A bulb is defined as a short, often plate-like stem Anatomical Concepts (Leaf and Stem) through which gas exchange takes place. The sto-
bearing a number of thick, fleshy leaves or leaf mata may be surrounded by normal epidermal
bases, which store water and nutrients; these are cells and are then anomocytic, but frequently they
bulb scales. The bulb scales may vary from one The vascular tissue is mainly in the form of pri- are surrounded by two or more cells differing in
to many, as in the Liliaceae s.str. mary vascular strands that consist of xylem (with size and shape from other epidermal cells; these
(Pseudobulbs, as found in many orchids, are di- tracheids and/or vessels, see below), and phloem, are called subsidiary cells. A stoma and its subsidi-
lated parts of the aerial stem that store water and which consists mainly of sieve tubes and their com- ary cells make up the stomatal complex. When
nutrients. They are corms rather than bulbs.) panion cells. there are two subsidiary cells, one alongside each
Velamen is a water-storing tissue in the outer Although the vascular strands in monocotyledons guard cell, as in many monocotyledons, the stoma
layers of some roots. It has a parchment-like ap- initially arise in a single ring, secondary bundles is paracytic; when there are four or six subsidiary
pearance and consists of one to several layers of soon develop so that the strands appear ““scat- cells surrounding the guard cells the stoma is tetra-
non-living cells with thickened and lamellate cell tered”. No cambium is formed between the cytic or hexacytic respectively. As will be explained
walls. Velamen is particularly common in epi- phloem and xylem where it is present in most dico- in the chapter on Distribution of Character Condi-
phytic orchids and aroids, but is also found in tyledons. Such an organization of vascular strands tions the ontogeny, i.e. the individual development,
the roots of many other monocotyledons, in par- is called an atactostele, and the condition atacto- of the stoma and the origin of the subsidiary cells
ticular in the Liliiflorae (see BARTHLOTT 1976a). stely. Tt is contrasted with the condition of eustely, is not always reflected in the appearance of the
ANCESTRAL
found in dicotyledons. There, a single ring of pri- mature stomatal complex; a special terminology
COMPLEX
mary vascular strands is formed. Then a cambium covering this situation is given in that chapter.
develops within and between strands, producing Trichomes are processes arising from the epidermis
The Aerial Stem Fig. 1. Sieve tube plastids in monocotyledons and their (hairs etc.). The hairs may be unicellular, i.e. com-
xylem on the inside and phloem on the outside, presumed evolution, according to BEHNKE (1981a). The
so that a cylinder of secondary tissue is produced, forms with cuneate protein crystalloids are called PIIc prise extensions of epidermal cells, but more often
Branching is monopodial when the shoot grows ap- : an eustele. Where secondary tissue occurs in (=cuneate). Protein filaments (f) are found in certain consist of a single row of cells. Sometimes, as in
monocotyledons a meristematic tissue produces monocotyledons (PIl¢f) and in some there are starch Bromeliaceae (Fig. 1541), the trichomes have a
ically, the main shoot generally exceeding the later- grains (s) as well as the protein crystalloids (PIIcs).
al ones in length. It is sympodial when axillary new sets of isolated vascular strands outside those short row of cells at the base and are branched
Sometimes other protein crystalloids (¢’) are found be-
branches successively take over the growth, as first produced. side the normal cuneate ones. above to form a stellate (star-shaped) or peltate
when the main shoot develops into an 'inflores- The xplem consists of tracheids or tracheids and (shicld-shaped) head. Some peltate hairs are of im-
cence or tendril, or dies off. A sympodium is a vessels. The former are elongate living cells, the portance for water uptake. Multicellular hairs with
sequence of such lateral shoots which successively cavities of which are without direct contact with forms with starch and filamentous protein respec- a broad multicellular base occur on the leaf mar-
overtake their predecessors (Fig. 52A). one another. Vessels begin as living cells which tively (Fig. 1). gins of Luzula (Juncaceae). Microhairs are small,
In most monocotyledons the acrial stem is herba- are produced in continuous rows; they die, and Laticifers are tubes or rows of elongate cells, con- generally bicellular thin-walled hairs found mainly
ceous, i.e. soft, usually green, and withering within acquire direct connections by means of perfora- taining a fluid of a somewhat milky appearance, in grasses (Fig. 1931-0). Glandular hairs are hairs
a limited time; in other groups it becomes tions in the end walls, the perforation plates. The in the monocotyledons only rarely coloured (as where one or more cells, generally at the end of
strengthened with lignin fibres and may be pro- end walls in narrow vessels are oblique and have in Dilatris of Haemodoraceae). the hair, are enlarged and secretory.
vided with bark. In some groups it becomes con- a row of transverse, narrow, slit-like perforations, Crystals of calcium oxalate occur in monocotyle- Intravaginal squamules, here interpreted as tri-
spicuously thick and long-lived, as in palms and separated by “ bars”, so-called scalariform perfora- dons rather frequently, most commonly as ra- chomes, are non-vascularized multicellular pro-
pandans, neither of which, however, has secondary tion; but wider vessels, when present, have a less phides, i.e. bundles of thin crystalline needles cesses situated in the leaf axils. They generally sec-
thickening. In other groups the stems are lignified oblique, or even transverse, perforation plate with (Fig. 28), which are contained in cells filled with rete mucilage, which is thought to protect the axils
but slender, and these plants are genuine shrubs; only a few perforations or one simple, circular per- mucilage. These cells may be almost isodiametric, from micro-organisms (TOMLINSON 1982).
they are common among the berry-fruited Aspara- foration. Roots, rhizomes, aerial stems and leaves but sometimes are elongated (“ sacs”) or form nar- Epicuticular wax is wax secreted on the outer sur-
gales. frequently differ in the type of xylem which they row tubes (“ raphide vessels™). In some groups oxa- face of the epidermis. It is frequently sculptured
In only a few families does secondary thickening contain. late occurs in the form of thicker solitary needles, in different ways, typical of the main groups of
occur, and this is of a kind different from that The sieve tubes of the phloem contain small plas- so-called styloids (pseudoraphides), which occur in monocotyledons (BARTHLOTT and FROHLICH
found in dicotyledons, as explained on p. 45. Most tids (leucoplasts). These may store starch and/or suberized (cork) cells. These are known in Ponte- 1983). See also the chapter on Distribution of
of these plants have a thick woody trunk. protein. The occurrence of protein [as compact deriaceac and Philydraceae, where raphides are Character Conditions.
The stems in a great many groups are hairy (see bodies (crystalloids) or as an annular structure of rare, and in Nolinaceae, Phormiaceae and Agava-
below) but thorns and spines are rare, being found thin filaments] and starch characterizes different ceae, where raphides are lacking.
in Smilacaceae and Petermanniaceae. The stems groups of angiosperms. All monocotyledons have Silica is deposited in several major groups of
Leaves
are climbing (scandent) in both of these families a number of triangular (cuneate) protein crystal- monocotyledons, cither as numerous small gran-
as well as in Dioscoreaceae and some other groups. loids in their plastids; a few also have starch and ules (silica sand) or as larger bodies of various
some have filaments. According to BEHNKE (1981), sizes and shapes. These shapes are described on The arrangement of the leaves on the stem is called
who has studied these structures, the monocotyle- p. 63. The silica bodies are often deposited in spe- phyllotaxy. When three or more leaves are placed
donous sieve tube plastids are of the PIIc Type cial short epidermal cells, silica short cells. at the same level, they are said to be whorled (verti-
(P=protein, Il =type 2 according to BEHNKE, |. c., Stomata (sing. stoma) are epidermal structures cillate), when in pairs they are opposite and when
c=cuneate), with PIlcs and PlIlcf representing composed of two guard cells embracing a pore solitary alternate. Where successive pairs of oppo-
4 Morphological Concepts The Inflorescence 5

are serrate, or beset with spines (Aloé, Agave, many The Inflorescence
genera of Bromeliaceae).
‘When distichous, the leaves, or their basal parts
only, are sometimes strongly compressed from the Inflorescences are floriferous branch systems more
sides and their adaxial surface (i.e. that facing the or less distinctly delimited from the vegetative part
stem) merely forms a groove basally which acts of the plant. Inflorescences can be divided into
sheath as a sheath and is obliterated above. Such leaves determinate (closed), where the primary axis is ter-
are described as ensiform or unifacial, in contrast minated by a flower, and indeterminate (open),
to the horizontally flat and bifacial normal leaves. where the axis does not terminate in a flower.
They occur in several families, e.g. Iridaceae, Or- Flowers of determinate inflorescences are nor-
open sheath closed sheath chidaceae and Haemodoraceae. mally actinomorphic, whereas indeterminate inflo-
In a few taxa the leaves are twisted through 180° rescences can have either actinomorphic or zygo-
Fig. 2. Terms used for the bases of (chiefly grass) leaves at the base so that the morphologically abaxial morphic (or even asymmetric) flowers (see below,
(above) and for a palm leaf (below). (Orig. B. JOHNSEN)
side is adaxial; we refer to this condition as “re- Floral Symmetry).
versed leaf blades™ (it is sometimes called resupin- Determinate inflorescences may be divided into
ate). panicles and cymes. In panicles there are lateral
The terminology of the shape of the leaves follows flowers, and branches terminated by flowers, at
(adaxial) hastula Potamogetonaceae. The homology of stipule-like general botanical practice. several levels below the terminal flower (as in Tri-
structures is not in every case established, for ex- Ptyxis is a term that refers to the folding or rolling cyrtis). Panicles may be many-flowered and com-
ample, hyaline extensions and stipular lobes (as of individual leaves in the bud stage. Types of plex or few-flowered. The panicle concept is also
in Joinvillea and Juncus species) which are always ptyxis in monocotyledons are the conduplicate, used for grasses, where the flowers are substituted
pseudopetiole lateral on the leaf near its base. A ligule is a hyaline conduplicate-plicate, plicate, involute, supervolute, by spikelets (which are indeterminate units). Pani-
extension of the leaf sheath on the adaxial side and explicative. (Ptyxis is sometimes included in cles may be distally expanded and more or less
of the leaf; it is typical of most grasses but is also vernation, a term which covers the arrangement flat-topped. Where the branches are very dense
known in Pontederiaceae. A contraligule is an ex- of parts in a bud with respect to each other). and the pedicels long this type may be umbel-
tension of the leaf sheath on the side opposite the Venation in monocotyledons is generally acrodro- like.
lamina. It occurs in the palms, grasses (Fig. 2) and mous, i.e. with longitudinal parallel or arched In cymes, lateral flowers are borne at only one
sedges. The hastula found in palms is at the distal veins, converging at the apex, whereas the veins level below the terminal flower, their stalks (pedi-
end of the pseudopetiole, but somewhat resembles in many of the primitive dicotyledons are campto- cels) arising in the axils of prophylls (bracteoles)
the ligule of the grasses. (See Fig. 2.) dromous-brochidodromous, with a midvein and, on the pedicel of the terminal flower. This process
A true petiole is present only in certain groups arising from it, lateral veins each of which is can be repeated indefinitely because every pedicel
site leaves are set at an angle of 90º to their prede- of monocotyledons. In other groups the leaves are arched to meet the one arising next above it (see has its own prophyli(s). The number of flowers
cessors the leaves are called decussate. The points linear and there is no differentiation at all between Hickey and WOLFE 1975). In leaves with a broad arising at the same level greatly affects the appear-
of attachment of successive solitary (alternate) petiole and lamina. Some regard such leaves as blade a minor, reticulate, vein system is interca- ance of the inflorescence; in the cymes of mono-
leaves form a spiral, the angle between successive phyllodial, consisting of a flattened petiole only, lated between the main veins. cotyledons there is generally only one, which
leaves often being constant. When the angle is but it may be more correct to interpret them as Cataphylls are scale-like leaves similar to the makes the cyme a monochasium. As the prophyll
180º, i.e. two leaves to a revolution, they are disti- the result of a particular mode of development sheath of a foliage leaf. They may be present at subtending it is generally on that side of the pedicel
chous (standing in two rows), and when it is 120º, of the leaf meristem. However, even these leaves the base of the shoot and as the first leaves (scales) which faces the preceding flower (the adaxial side)
i.e. three to a revolution, fristichous (standing in are frequently differentiated secondarily into a of lateral branches, especially those of the inflores- the cyme forms a zigzag and this is called a rhipi-
three rows). narrower proximal part, a pseudo-petiole, and a cence. dium. Where the prophyll and its axillary flower
The leaves of some monocotyledons, e.g. Dios- broader distal part. In certain bamboos and other The shoot may continue into an inflorescence, or are not strictly adaxial the monochasium may re-
corea, are differentiated into a short leaf base, a grasses and in palms, the pseudo-petiole may be the inflorescence may be distinctly separated from semble a bostryx (helicoid cyme). A bostryx in the
distinct petiole and a blade (lamina). However, the long and clearly delimited. the vegetative part, especially where there is a leaf strict sense results when each pedicel arises in the
leaf base more often encloses the stem completely When the leaf is differentiated into a distinct pe- rosette, and the inflorescences are borne each on axil of a lateral prophyll (i.e. one set at right angles
and forms a sheath of variable length which can tiole and lamina, as in some Dioscoreales and Ar- a leafless peduncle. A peduncle arising from a basal to the preceding one) and the direction of rotation
be open, the two margins of the sheath being free, ales, the lamina may be lobed or even compound. rosette is known as a scape and the plant (or inflo- is always the same. The umbel-like inflorescence
or closed, when the margins are fused with each Pseudo-compound leaves are leaves which when ini- rescence) is described as scapose, a condition found of Alliaceae and Amaryllidaceae is probably de-
other. Regardless of whether it is sheathing or not, tiated are simple, but which in the course of devel- in Hyacinthaceae and Amaryllidaceae, for exam- rived, by condensation, from this type of cyme.
the leaf base may be extended into lateral lobes opment split along the nerves to become “com- ple. Sometimes, each new flower emerges in the axil
or stipules. The base of a linear leaf, or leaf blade pound”, as in many palms and Cyclanthales. In of an abaxially (opposite to adaxially) placed brac-
in the case of a leaf with a sheath, may be pro- Musaceae and some other families, simple leaves teole, resulting in a drepanium, a rare type of inflo-
longed on either side of the stem as a lobe, the may become torn so as to appear compound. The rescence (in some Juncaceae).
auricle (ear). The stipules may be displaced into leaf margins in monocotyledons are typically en- Indeterminate inflorescences may be divided into
the leaf axil to form a stipular sheath, as in many tire, but in some, mostly succulent species they thyrses, where the pedicels of lateral flowers bear
 6 Morphological Concepts The Perianth 7

new flowers in the axils of their bracteoles (these arate plants, or monoecious, with both kinds of The Floral Axis The Perianth
lateral components are thus lateral cymes), and flowers occurring on the same plant. Sterile
racemose inflorescences (botrya) where this is not (neuter) flowers lack both functional stamens and
The floral axis (receptacle) in monocotyledons (see The perianth (perigone) represents the floral enve-
“ so. The latter are classified according to the length functional pistil(s); such flowers are often special-
above, The Flower) is generally not strongly devel- lope and consists of floral (perigonal) leaves (te-
and thickness of the main axis and the length of ized for the attraction of pollinators.
oped. Thus a floral disc, i.e. a disc-shaped or annu- pals). In most monocotyledons the perianth con-
the pedicels: racemes have a long inflorescence axis In connection with the symmetry of the flower
lar process developed from the receptacle, is ex- sists of two whorls of tepals which are either simi-
and long pedicels, umbels have a short inflores- the concepts of the median and transverse planes
tremely rare, whereas in dicotyledons this structure lar or dissimilar. Even when dissimilar they are
cence axis and long pedicels, spikes have a long, are used. The median plane of a lateral flower is
is common and often functions as a nectary. In often not readily divisible into outer green sepals,
slender inflorescence axis and very short pedicels that falling through both the inflorescence axis and
dicotyledons the receptacle is also frequently ur- and inner contrastingly coloured petals, though
or none, spadices (sing. spadix) have a long fleshy the pedicel and main axis of the flower. The trans-
ceolate, but urceolate structures of monocotyledon this is so in some groups (taxa of Alismatales, Bro-
inflorescence axis and very short pedicels or none verse plane of a lateral flower is that cutting the
flowers are mainly formed by the fused tepals (and meliales, Commelinales, etc.). When green and se-
and capitula (heads) have a short inflorescence floral axis at right angles to the median plane.
stamens). See below under perigyny. pal-like the tepals are described as sepaloid, and
axis, and very short pedicels or none. Neither of these terms can be applied to a flower
when of a colour other than green (white or bright
The inflorescences of many monocotyledons are that is terminal on the inflorescence axis.
colours) as petaloid. In dicotyledons the sepals are
extremely complicated, and painstaking analyses A flower is actinomorphic (polysymmetric, radially
often collectively termed the calyx and the petals
may be necessary to reveal their true nature. Thus symmetric, “‘regular ) when three or more planes Numerical Conditions and Insertion
the corolla, and this is also possible in monocotyle-
the superficially simple “spike” or “spadix” of of symmetry (giving mirror images) can be placed of the Components of the Flower dons showing this kind of differentiation, but we
Typha and the “head” of Sparganium have proved through it. It is bisymmetric when two planes of
shall not follow this usage here.
to be complex, branched inflorescences (D. symmetry can be placed through it, and zygo-
A flower is described as cyclic when all the organs When the tepals in the two whorls differ conspic-
MULLER-DOBLIES 1968 and U. MULLER-DOBLIES morphic (monosymmetric) when only one plane of
of the same type are in whorls (for whorled, see uously the perianth is termed heterochlamy-
1969, respectively). The asymmetric flowers of symmetry can be placed through it. The zygo-
above, Leaves). This is the condition in perhaps deous.
Marantaceae are aggregated into complex thyrses morphic flowers of monocotyledons always have
all monocotyledons, whereas in many dicotyledons Flowers in which the tepals are fused are called
(ANDERSSON 1976, 1981), and the dense ““fascicles” a median plane of symmetry. The twisting of the
all or some of the floral parts are spirally set, the syntepalous, and the condition syntepaly. Tepals
of Bobartia, Iridaceae, into complex panicles pedicel or ovary through 180°, inverting the flow-
flowers being acyclic or hemicyclic respectively. are often fused to form relatively narrow floral
(DAHLGREN, unpublished). er, is called resupination; it occurs in several
The number of whorls of floral parts (prophylls tubes. The tepals may taper basally into a stalk
groups and is prevalent in orchids. Asymmetric (ir-
excluded) in the flower is indicated by the terms and are then described as clawed (unguiculate), a
regular) flowers have no planes of symmetry at
pentacyclic (with five whorls, which is a common rare condition in monocotyledons.
all (examples: Cannaceae, Marantaceae).
and probably ancestral state in monocotyledons), Some or all of the tepals may be provided basally
When describing a lateral flower, the lower side,
The Flower tetracyclic (with four whorls; as when one whorl with a nectary known as a perigonal nectary. The
which faces away from the inflorescence axis, is
of stamens is lacking), tricyclic (with three whorls), nectarial area is sometimes recessed to form a
described as abaxial, and the upper, facing to-
wards the inflorescence axis, as adaxial. These
ete. pouch or spur (Fig. 108 H and N).
The flower is situated in the axil of a subtending
A whorl of floral parts is classified according to A median labellum, or lip petal, is present in some
leaf or bract. It consists of a pedicel (flower-stalk), terms are also used to describe respectively the
its number of components (merism, merous condi- families. This is so in Orchidaceae, where it is the
which usually bears a prophyll (bracteole) on the outer and inner sides of floral parts in relation
tion). Thus nearly all monocotyledons are trimer- median, upper tepal of the inner whorl (though
adaxial side (i.e. the side opposite the subtending to the floral axis (i.e. the centre of the flower).
ous (parts in threes), but some are dimerous (parts the flowers are usually resupinated), and in
leaf and towards the parent axis). In the monocot- Floral diagrams are constructed to illustrate and
Zingiberaceae and Costaceae, where it consists of
in twos), or fetramerous (parts in fours).
yledons the prophyll is frequently two-ribbed, sug- compare, in a uniform and schematic manner, the
According to the position of the tepals and sta- the two lower petaloid staminodes of the inner
gesting that it may have arisen by concrescence (transverse) plans of flowers. The floral compo-
mens in relation to the ovary of the pistil (or the staminal whorl fused together. Though non-ho-
of two lateral prophylls (such as are present in nents are placed in the diagram in such an order
pistils), distinction is made between hypogynous mologous, the labellum in each group has a similar
most dicotyledons). Flower buds may also develop that the lowest and/or outermost parts are on the
flowers, where tepals and stamens arise from the function, namely as a landing place for pollen vec-
in the axil of the prophyll, as in cymose inflores- periphery and the uppermost/inner parts are in the
floral axis “ below the gynoecium ™, and epigynous tors.
cences. Rarely two or more prophylls are present centre (i.e. in the order bracteoles-tepals-stamens-
flowers, where these parts arise above the ovary, When tepals are lacking the flowers are described
on the pedicel. pistils). It is usual to employ standard symbols
““on the gynoecium”, their basal parts being then as atepalous (naked) or, when the loss of the tepals
The pedicel terminates in the floral axis (recepta- for homologous parts. Empirical floral diagrams
is obvious, apochlamydeous.
fused with the pistil wall, taking part in the forma-
cle), on which the tepals (perianth), stamens (an- are those in which the components of the flower
tion of the wall of the ovary and fruit. More rarely The paracorolla (corona) is a structure derived
droecium) and pistil(s) (gynoecium) are inserted. are shown in their position without any attempt
the flowers are hemi-epigynous, having the tepals from appendages of stamens or tepals as, for ex-
The flower is termed complete when perianth, sta- at interpretation, whereas theoretical floral dia-
and stamens inserted halfway up the ovary. Peri- ample, in Amaryllidaceae and Velloziaceae.
mens and pistil(s) are all present, incomplete if any grams involve an interpretation, for instance they
gynous flowers, where there is a cup-shaped dila-
of these parts are missing. It may be bisexual (per- may indicate supposedly lost parts by crosses, and
tion of the receptacle, free from but surrounding
fect, hermaphrodite), having both stamens and divisions and fusions by other suitable symbols.
the ovary or part of it, are common in some groups
pistil(s), or unisexual (imperfect), when it lacks ei-
of dicotyledons but are strictly speaking not found
ther stamens or pistil(s). Species with unisexual
in monocotyledons.
flowers may be cither dioecious, with pistillate (fe-
male) flowers and staminate (male) flowers on sep-
 The Anther Wall and Tapetum 9
8 Morphological Concepts

The Androecium According to the position of the microsporangia

in relation to the connective, the anthers can be
Androecium is the collective term for the stamens divided into introrse, facing towards the centre of
and stamen homologues (i.e. also staminodes, see the flower, extrorse, facing away from the centre
below). By definition, stamens are the floral struc- of the flower, and latrorse, facing laterally. Gener-
tures that carry microsporangia in which the micro- ally these are also the directions in which the mi-
spores are formed and subsequently develop into crosporangia empty their pollen, which they do
pollen grains. through longitudinal slits. In some groups, how-
Diplostemonous flowers, where two whorls of sta- ever, they dehisce by one, two or four apical pores
mens are present (diplostemony), are here consid- and are then called poricidal; in these it is still
ered to be the ancestral state in the monocotyle- possible to determine whether they are introrse or
dons. Flowers with only one whorl of stamens are extrorse. When the anthers dehisce by longitudinal
haplostemonous, the condition, haplostemony, be- slits, the wall separating the two microsporangia
ing assumed to have resulted from loss of either of each theca has generally broken down before-
the outer or the inner staminal whorl. hand, so that only a single line of splitting is re-
Stamens are numerous in several groups of mono- quired to open each theca.
cotyledons, presumably by secondary multiplica- In some monocotyledon flowers the filaments are
tion (“dédoublement”), and the androecium is laterally fused (connate) at the base to form a sta-

oe
then described as multistaminate or pleiomerous. minal tube. In male flowers of Araceae the stamens
The stamens consist of a generally slender stalk, are often totally fused to form a synandrium; the
the filament, and an anther. The anther consists fusion of the anthers is termed synanthery. Rarely,
of the connective, which is the continuation of the the filaments are fused with the style to form a
filament, and two thecae, each of which consists gynostemium (column).
of two microsporangia (locules, pollen sacs). Rare- The terminology of the numerous specializations
ly, by reduction, the anthers have only one theca in the flowers of Orchidaceae appears under that
and are called monothecous, or have thecae that family (pp. 255-259 and Figs. 119, 122, 124, 125,
consist of only one microsporangium which are 126).
called unisporangiate. Staminodes are sterile homologues of stamens and
In some monocotyledons, e.g. many Dioscoreales, may or may not have rudimentary anthers. Sta-
the stamens are flat and somewhat leaf-like, and minodes are considered to be derived (in the phylo-
in some members of this order as well as many genetic sense) from functional stamens, and sta-
others, the microsporangia are attached below the mens and staminodes together generally do not
exceed six in any flower. When they are flat and Fig. 3. Stamen characters. A and B comparison between the inner parietal layer divides as in the Basic Type
apex, which is then often described as a connective hypopeltate (A) and epipeltate (B) anthers, the asterisk and the outer becomes the endothecium directly,
tip or appendage. In these kinds of stamens the brightly coloured, as in many Zingiberales, they (%) representing the morphological upper side. C “ Un- so that again there is only one middle layer. In
anthers are not clearly set off from the filament are known as petaloid staminodes (Fig. 169H). differentiated” anther. D Basifixed, impeltate, non-sagit-
tate anther. E Basifixed, impeltate, sagittate anther. the Reduced Type, finally, neither parietal layer
and may be described as “undifferentiated”
F Basifixed, impeltate, x-shaped anther. G-J Sequence divides and there is no middle layer; the outer
(ScHAEPPI 1931). Where the anther is attached at
to explain derivation of adnate (H), semiadnate (1) and forms the endothecium directly and the inner the
its base to the filament it is described as basifixed
The Anther Wall and Tapetum dorsifixed (J) anther types from an initial (G). K “ Undif- tapetum. This type is extremely rare in dicotyle-
(impeltate). Basifixed anthers are called sagittate ferentiated ” anther, exemplified by Sparganium. L Series
of transverse sections through an undifferentiated dons, and in monocotyledons is known only in
when the thecae are lobate and divergent from
The layers of the microsporangium wall can be anther, exemplified by Smilax. M-N Stamens of Yucca Najas (Najadaceae) and in Lemna and Wolffia
the connective at the base; otherwise they are non-
sagittate. The anthers are described as dorsifixed classified according to the behaviour of the two Sfilamentosa, M fully developed in lateral view, N in juve- (Lemnaceae).
nile stage, front view. O Anther of Dianella caerulea, The endothecium is a layer in which the cells are
(peltate) where the anther with its connective ex- parietal cell layers which line the microsporangial with filament dilated below anther. (All after SCHAEPPI
epidermis and invest the microsporogenous tissue. provided with wall thickenings which play a part
tends below its point of attachment, which is thus 1939; from WEBERLING 1981)
There are four different types, named Basic, Di- in the dehiscence of the microsporangial walls. The
located somewhere along the mid-line of the
cotyledonous, Monocotyledonous and Reduced wall thickenings (DAHLGREN and CLIFFORD 1982)
anther. Peltate anthers can be divided into epipel-
can be divided into two general types, the Spiral
tate when the part of the anther that is prolonged (Davis 1966).
downwards beyond the attachment point of the The Basic Type is not known in monocotyledons In the Dicotyledonous Type, which is known and the Girdle Types.
and is rare in dicotyledons; it involves one peri- among the monocotyledons only in Tacca (?), but The tapetal layer (tapetum) is best developed when
filament faces inwards, and hypopeltate when this
clinal division of each parietal layer to form four is common in dicotyledons, the outer parietal layer the microspores are in the tetrad stage. The tape-
part faces outwards in relation to the centre of
layers in all: an outer layer which becomes the divides as in the Basic Type, but the inner becomes tum then surrounds the spores and supplies them
the flower. These two types have somewhat differ-
ent distributions, the former being common in Li- endothecial layer with characteristic wall thicken- the tapetal layer directly, so that there is only one with nutrients. There are three types of tapetum
ings (see below), two middle layers and an inner middle layer. In the Monocotyledonous Type, (1) secretory or glandular, in which the tapetal cells
liales, the latter in Asparagales (HUBER 1969). (See
which is by far the commonest in monocotyledons, remain in their initial position but lose their walls
Fig. 3.) layer which gives rise to the tapetum (see below).
10 — Morphological Concepts The Gynoecium 11

and finally degenerate in situ, (2) amoeboid, in dominant. The Successive Type is sometimes sulculate, where two (rarely three) slit-like aper- The number of carpels in the gynoecium is indi-
known as the Monocotyledonous Type. tures are present and lie in the equatorial plane cated by the terms monocarpellary, bicarpellary,
which the tapetal cells persist for some time but
eventually form a periplasmodium and invade the The terminology of pollen morphology (Fig. 4) of the pollen grain; tricarpellary, or multicarpellary (with one, two,
cavity occupied by the microspores, and (3) peri- here follows ERDTMAN (1952) as in DAHLGREN and zonisulculate or meridionosulcate, where there is three or many carpels, respectively), conditions
plasmodial, where the cells dissolve at an early CLIFFORD (1982). The pollen grains of monocotyle- one continuous aperture all round the equator of which all occur in the monocotyledons.
stage to form a periplasmodium. The last two dons generally have only one aperture (a thin part the pollen grain, thought to have arisen through The pistils, whether monocarpellary or consisting
types are often known collectively as the amoeboid- or hole in the exine, which is the outer layer of fusion of two sulculi, and of two or more carpels, may comprise the follow-
the pollen grain wall). The aperture can be located porate, where one or more round apertures are ing parts:
periplasmodial type (Fig. 32).
at the distal pole, i.e. the part of the pollen grain present in the equatorial plane of the pollen grain a stipe, which is the stalk of the pistil; it is generally
that is directed away from the centre of the pollen (this in its typical form is not found in monocotyle- very short but may in exceptional cases be long,
tetrad. When a distal aperture has the shape of dons but see, however, Carludovica in Cyclantha- as in Ruppia (Potamogetonaceae) ;
ceae). an ovary, the widened part enclosing one or more
The Pollen Grains and Their Dispersal a slit (sulcus) the pollen grain is called (mono-)sul-
cate and when it has the shape of a circular hole Colpate pollen grains, where slit-like apertures cavities (locules) containing the ovules and charac-
(ulcus) it is called wulcerate. These two types are (colpi) are present in the equatorial plane and di- terized as unilocular, bilocular or trilocular accord-
The interior of the microsporangium is occupied the commonest in the monocotyledons. In other rected at right angles to it, are probably not found ing to the number of locules (ovaries containing
by the archesporial cells, which give rise to the cases the exine is very thin or not even coherent, in the monocotyledons. more than three locules are rare in monocotyle-
microspore mother cells ; these undergo meiosis, in- as in the Najadales of the Alismatiflorae, the The pollen is generally dispersed as separate grains dons); and
volving two successive divisions, to produce the Triuridiflorae, most Zingiberiflorae and most taxa but occasionally it is coherent in tetrads or massu- a style, the narrow upward prolongation of the
microspore tetrads. This meiotic division is known of Orchidaceae of the Liliiflorae, in which case lae. In the mature pollen grain division of the mi- ovary carrying the stigmatic papillae; the carpels
an aperture is naturally not visible, and the pollen crospore cell has taken place, so that either the of a pistil, when more than one, may each have
as microsporogenesis and it proceeds in one or
other of two modes, the Successive and the Simul- grain is called inaperturate, although this is prob- pollen grain contains two cells, the vegetative and a separate “style” which is then called a stylodium,
taneous (Fig. 33). In the Successive Type a cell wall ably inaccurate in most cases and the grain is bet- the generative cell, or, where the latter has divided or they may share a single common style (style
is formed after the first division, and another in ter described as omniaperturate (i.e. with the aper- further, as happens in certain groups, it contains in the strict sense) and this may be three-branched
ture covering the whole pollen grain) (THANIKAI- three cells, the vegetative cell and two sperm (tribrachiate), three-lobed (trilobate), or entire,
each of the daughter cells after the second meiotic
division; the cells of the tetrad usually lie in one MoONI, 1978). Some aperture conditions that are cells. when it may end in a head (being capitate) or a
rare or not definitely known in monocotyledons The pollen may be dispersed by animals (zoo- point (being punctiform); the style is generally ter-
plane and the arrangement is square, T-shaped or
linear. In the Simultaneous Type the two perpen- follow: gamy), wind (anemogamy) or water (hydrogamy). minal, i.e. situated on the top of the ovary, but
trichotomosulcate, where the sulcus is trilobate; Dispersal by insects (entomogamy), birds (ornitho- in some groups it emerges from the side or even
dicular nuclear divisions proceed before wall for-
mation starts, and wall formation occurs by the spiraperturate, where one or more furrow-like ap- gamy) and bats (chiropterogamy) are the most im- the base of the ovary (gynobasic), as in many Al-
ertures are spirally arranged on the pollen surface; portant types of zoogamy. Flowers show many ismataceae and Triuridaceae; the stylodia may be
advance of constricting furrows from the peri-
phery of the tetrad, which is tetrahedral. Both foraminate, where a number of round apertures specializations for these types of pollination. The bifurcate nearly to the base, as in some Najadales
(foramina) are evenly distributed on the pollen sur- list of specializations which characterize a particu- (Fig. 150D) and some Iridaceae; when there are
types are widespread in the monocotyledons,
face; lar method of pollination, some or all of which stylar lobes, these are generally opposite the mid-
whereas in dicotyledons the Simultaneous Type is
are expressed in all plants adapted to that method, vein of the carpel;
is termed a pollination syndrome, for example wind stigmatic papillae, cells receptive to pollen tubes
pollination is characterized by the “syndrome of and situated on the distal parts of the stylodia,
anemogamy™. Some monocotyledons exhibit hyd- the style or its branches or lobes, being either re-
rogamy (water pollination) and in the more ad- stricted to their actual apex or, especially in wind-
A ' B !: C ' D vanced marine exemplars of this method the pollen pollinated taxa, covering a considerable part of
grains are almost completely devoid of exine and, their surface.
di. po. 7 © T lWO [) although globose at first, they rapidly elongate to Monocarpellary pistils in the monocotyledons al-

%º
ways have a unilocular ovary, whereas the bi- and

Moo=l
become filiform, in which condition they are more
o easily caught by the stigmas. This elongation may tricarpellary pistils generally have two and three
be equivalent to the development of a pollen tube separate locules respeetively, the locules being sep-

VUV 1Ã
/o - in other groups. arated by partitions (septa). In several groups,
however, the syncarpous pistils lack septa, or have
incomplete septa, so that they are in effect unilocu-
prpo.L' o < W O lar.
The Gynoecium The placenta is defined as that part of the ovary
1 2 3 4 5 6 7 8 on which the ovules are borne. When the ovules
ig. 4. Terminology of pollen grain apertures. Abbrevia- ertures, which are either slit-like or circular, are classified Gynoecium is a collective term for the carpels of are numerous the placenta is a richly vascularized
tions: di.po. distal pole; equ. equator; pr.po. proximal accordingly: (1 laesura); (2 hilum); 3 sulcus; 4 ulcus; a flower, which can be free and make up separate tissue. The placenta is generally confined to a par-
pole; I apertures polar; Il apertures non-polar; IA aper- (5 colpi); (6 pori); (7 rugae); & foramina. Those placed pistils, a condition called apocarpy, or fused into ticular part of the locule (base, apex or central
tures in proximal pole; IB apertures in distal pole; in parentheses not found in monocotyledons. (After
IIC apertures equatorial; IID apertures global. The ap- ERDTMAN 1952) a single pistil, a condition known as syncarpy. axis of the pistil).
12 — Morphological Concepts The Ovule

When there is a single carpel placentation can be cotyledon family constantly has only one. Ovules
classified as: with two integuments are termed bitegmic, those
marginal, with the ovules inserted along the carpel with one integument are unitegmic and those in
margin, which the integuments have been lost (rare) are
apical, with the ovules concentrated at the top of called ategmic. The chalaza (chalazal part) of the
the locule, ovular body is generally that located at the end
basal, with the ovules concentrated at the bottom opposite to the micropyle.
of the locule, and In addition, tissues that develop into arils, stro-
laminar (-dispersed), with the ovules scattered over phioles, caruncles and analogous structures are
the whole or most of the inner wall of the carpel. present in scattered groups (see below under Seed).
In syncarpous pistils the placentation is classified Arils, when present, may form a conspicuous enve-
according to the position of the placentae in the lope outside the outer integument as early as the
ovary as follows: ovular stage, e.g. in Asphodelaceae (Fig. 5S-T)
parietal, with the ovules inserted on the wall of and many Zingiberales.
the ovary along the carpel margins, which is gener- According to the number of layers of cells sur-
ally the case with unilocular ovaries (without rounding the archesporial cell and embryo sac (see
septa), below) the ovule is termed crassinucellate (several
central or axile, with the ovules on the central axis or numerous layers) or fenuinucellate (one or few
of the ovary, layers). Within the nucellar tissue one primary
free-central, with the ovules borne on a central archesporial cell (rarely more) will differentiate.
column or at least centrally at the bottom of a This may cut off one or more cells (parietal cells)
unilocular pistil (e.g. Luzula). at the micropylar end before the two meiotic divi-
By combining these two sets of concepts the pla- sions leading to the megaspore tetrad (cf. The Pol-
centation can be described adequately. Certain len Grains). The parietal cells, when present, gen-
trends are observable in monocotyledons; thus, erally divide further, forming parietal tissue be-
when the ovules are apical they tend to be ortho- tween the embryo sac (see below) and the nucellar
tropous (see below) and when the ovules are ex- epidermis. In a number of groups a parietal cell
tremely numerous the ovary is often unilocular is not cut off; this is often the case where the ovules
and the placentation parietal (Orchidaceae, many are tenuinucellate. Sometimes the nucellar epider-
Burmanniales, some Philydrales, Mayacaceae). mis in the apical part of the nucellus divides by
periclinical divisions to form what is known as
a nucellar cap. If this is the case in ovules without
parietal tissue some embryologists refer to such
The Ovule ovules as “pseudocrassinucellate” (here belong
many grasses, most Cyclanthaceae except Cyclan-
An ovule consists of the following parts: thus and a few other groups).
the funicle, the stalk by which the ovule is attached According to their morphology ovules are divided
to the placenta; in curved anatropous ovules the into the following types: Fig. 5. Embryological details as exemplified by Aloé ci-
liaris (Asphodelaceae). A-D Anther wall development
funicle is fused with one side of the ovule and anatropous, with a straight nucellar axis, and re- according to the Monocotyledonous type, showing for-
is visible as a rib or ridge, the raphe, flexed upon the elongated funicle, which is usually mation of three layers under the epidermis, the inner-
the nucellus, the central region of the ovule (often fused at the side to the ovular body (=raphe; see most becoming the tapetum (here of the secretory type)
regarded as being equivalent to a megasporan- above); the micropyle is thus close to the funi- and the outermost (apart from the epidermis) the en-
dothecium, the wall thickenings of which are here spiral.
gium), and cle; E-O Microsporogenesis, here of the simultaneous type
one or two integuments which are two or a few hemianatropous (hemitropous), with a straight nu- with two successive nuclear divisions and wall formation
cell layers thick and envelop the nucellus. cellar axis at right angles to the funicle; proceeding from the periphery inwards. The pollen
The micropyle is the more or less narrow, rarely campylotropous, with a curvature of the nucellar grains are sulcate. P-Q Transverse and longitudinal sec-
completely obstructed, pore at the top of the inte- axis, which brings the micropyle near the funicle tions of ovary showing axile placentation. R-T Develop-
ment of ovule, showing a third layer, outside the integu-
guments. The micropyle can be formed in both and makes it basally directed; the embryo sac in ment, which represents the aril so typical of the Aloé
integuments or in either the inner or outer integu- this case is more or less straight, but if the nucellar group in Asphodelaceae. U-W Formation of a parietal
ment, if one or the other integument is delayed axis is even more curved and the embryo sac is cell which divides later to form a parietal tissue.
in development. It may have a “zigzag” form if also curved the ovule is described as amphitropous; X-Y Megaspore tetrad with the three micropylar spores
degenerating, Y being a case where the two megaspores
the pore of the inner integument does not exactly orthotropous (atropous), with a straight nucellar are still large. Z-E' Development of embryo sac accord-
coincide with that of the outer. Nearly all mono- axis and the micropyle opposite the funicle. ing to the Polygonum Type, from the binucleate stage
cotyledons have two integuments, and no mono- to the mature embryo sac. (GOVINDAPPA 1955)
14 Morphological Concepts The Fruit 15

Embryo Sac Formation which is the result of a form of nuclear fusion This division gives rise to a basal (proximal) cell to be a double fruit of follicles (see below) belong-
of the three lower nuclei at the four-nucleate stage, and a terminal (distal) cell. Further classification ing to adjacent monocarpellary flowers.
The primary archesporial cell or the cell remaining followed by two successive divisions of this triploid is based on whether the terminal cell divides trans- The fruit consists of the fruit wall or pericarp and
after the parietal cells have been cut off (see nucleus. (Haploid means having one set of chro- versely or longitudinally. Longitudinal division the seeds. When the flower is epigynous, the wall
above), functions as the embryo sac mother cell; mosomes and triploid means having three, the nor- gives rise to the Onagrad and Asterad Types, while of the fruit inevitably incorporates accessory parts
this divides by the meiotic process (two nuclear mal condition of the vegetative cells being diploid, three main types arise from transverse division, derived from the perianth and androecium or from
divisions) to form four nuclei, the megaspore nu- with two sets.) the Caryophyllad, Solanad and Chenopodiad the receptacle, and its pericarp is therefore not
clei. Generally wall formation takes place after Further types, the Clintonia, Adoxa and Drusa Types. fully homologous with that of a fruit developing
each division, producing four megaspores, only Types, all tetrasporic, are rare and need not be The Asterad and Onagrad Types are common in in hypogynous flower.
one of which takes part in the formation of the defined here (see, however, DAHLGREN and CLIF- monocotyledons. They differ mainly in the role Monocarpellary fruits in monocotyledons are clas-
embryo sac (see below), which is then monosporic. FORD 1982). of the basal cell — in the Asterad Type the basal sified as follows:
Sometimes wall formation does not take place cell plays an important part in the formation of Jfollicle — dry, usually dehiscing along the carpellary
after the second division, so that two cells (dyads), the embryo, whereas with the Onagrad Type its margin;
cach with two megaspore nuclei, are produced; role in the formation of the embryo is insignificant. achene — small, hard, dry and indchiscent, with
Endosperm Formation
one of these forms the embryo sac, which is then one or a few seeds;
bisporic. Rarely, but especially in monocotyledons, The latter type is characteristic of most orchids monocarpellary drupe — pericarp fleshy on the out-
no cell walls are formed in connection with meiosis The angiosperms are characterized by what is and of Cyperales but is on the whole less common side and hard within; indehiscent and generally
and the cell containing all four megaspore nuclei known as double fertilization, where the nucleus than the Asterad Type. A special modification of one-seeded ;
develops into the embryo sac, which is tetra- of one of the two sperm cells fuses with the nucleus the Asterad Type in which the primary division monocarpellary berry — fleshy, indehiscent and
sporic. of the egg cell (forming the zygote), and the nucle- is laid down obliquely is found in the grasses. with one or several seeds.
The following main types of embryo sac occur in us of the other fuses with the central nucleus or In the Caryophyllad Type (Fig.135L) the basal Syncarpous fruits (which in monocotyledons rare-
monocotyledons (see DAHLGREN and CLIFFORD the polar nuclei. The zygote (diploid) divides to cell does not divide further but increases in size ly consist of more than three carpels) are classified
1982): form the embryo; the fusion product of the nucleus and forms a vesicular suspensor for the embryo as follows:
The Polygonum (Normal) Type (Fig. 5X-E’), a of the sperm cell and the central nucleus or polar which is formed wholly from the terminal cell and capsule — dry and dehiscent, generally with few
monosporic type in which the lowest (chalazal) nuclei (usually triploid) divides to form the endo- has a neck that varies in length. It is found in to numerous seeds. The capsules are loculicidal if
megaspore undergoes three divisions to form an sperm. all (7) Alismatiflorac and most Ariflorae, but oth- dehiscent along the midribs of the carpels, septici-
eight-nucleate embryo sac with one egg cell accom- Three main types of endosperm formation oc- erwise occurs only sporadically in monocotyle- dal when breaking open and dehiscing along the
panied by two synergids at the micropylar end and cur: dons. sutures of the septa, and septifrage when the outer
three antipodals at the chalazal end. The remaining cellular, where the successive divisions of the endo- The Solanad and Chenopodiad Types are only oc- walls are shed as valves leaving the septa and the
two nuclei, the polar nuclei, may fuse to form a sperm nuclei are quickly followed by the formation casionally reported in monocotyledons and will (usually axile) placentae;
central nucleus. This is the commonest and most of cell walls, therefore not be described here. pyxis (pyxidium, lid capsule) — dry, circumscissile,
widely distributed type in monocotyledons and helobial, where the first division of the primary i.e. splitting round the capsule so that the upper
also in dicotyledons. endosperm nucleus is followed by a wall division portion forms a lid;
The Allium Type (Fig. 501-P), a bisporic type, in giving rise to a larger micropylar and a smaller syncarpous nut or nutlet — dry, more or less hard,
The Fruit
which the lower chalazal dyad undergoes two divi- chalazal chamber or division; in the micropylar indehiscent, and with one or a few seeds ;
sions to form an eight-nucleate embryo sac similar chamber the succeeding nuclear divisions are fol- caryopsis — a syncarpous nutlet in which the peri-
in organization to the Polygonum Type. This is lowed by wall formation after a delay, whereas The fruit, like the ovary, may be formed by one carp and seed coat are fused (found in most
known in several groups of monocotyledons (e.g. those in the chalazal chamber may or may not or several carpels. Where carpels are more than grasses);
Alismataceae, most Trilliaceae, and various genera be followed by wall formation, and one, the whole set of monocarpellary fruits of the schizocarp — usually dry, splitting radially along
of Alliaceae, Hyacinthaceae, Amaryllidaceae, nuclear, where the successive divisions of the endo- flower developed from an apocarpous gynoecium the septa so that complete locules are detached
etc.). sperm nuclei are not accompanied by wall forma- may be validly compared with the single fruit de- (mericarps); these are generally nut-like. Schizo-
The Endymion Type, a bisporic type which differs tion, which occurs considerably later. veloped from a syncarpous gynoecium (the main carps are very rare in monocotyledons but are
from the Allium Type in that the upper (micropy- The endosperm types are of importance in the difference being the degree of fusion of the car- known in Cyanastraceae and Juncaginaceae;
lar) dyad develops into the embryo sac. This is study of monocotyledon phylogeny. pels). syncarpous drupe — like the monocarpellary drupe
less common than the Allium Type and is found A distinction is usually made between an aggregate but consisting of two or more carpels;
in a few Convallariaceae, Hyacinthaceae, etc. Sfruit, which is composed of a number of coherent syncarpous berry — like the monocarpellary berry,
The Fritillaria Type (Fig. 112M-T), a tetrasporic individual fruits formed by separate pistils in the but consisting of two or more carpels.
Embryo Formation The utricle should not be equated with the forego-
cight-nucleate type, with two divisions following same flower (virtually absent in monocotyledons)
upon meiosis but accompanied in a characteristic and multiple fruits, which are a group of coherent ing fruits; it consists of a nutlet which is tightly
way by nuclear fusions. The nuclei of the egg cell Embryo formation (embryogeny, embryogenesis) is fruits formed from adjacent flowers (as in Ananas, enclosed in a flask- or bladder-like envelope and
and synergids and the upper polar nucleus are hap- classified according to the planes of division of Cyclanthus and Carludovica). This distinction is is known in Carex and some other Cyperaceae.
loid, and are all derived from the micropylar me- the zygote (see above, Endosperm Formation). lost when both of these are indiscriminately called The term is also widely applied to the thin-walled
gaspore nucleus. The three antipodals and the The first division of this cell is generally transverse, syncarpia. The capsule-like fruit of Gaimardia in fruits of some grasses, in particular Eleusine and
lower polar nucleus are triploid instead of haploid, and this is the case in all known monocotyledons. Centrolepidaceae (Fig. 215M) is here considered its allies.
16 Morphological Concepts

The Seed and in the Hydatellales. Chalazosperm is a storage

tissue that consists of proliferating tissue from the Chemical Characters
The seed consists of : chalazal part of the ovule. It occurs together with
the seed coat, which is formed from the integu- perisperm in Costaceae, Cannaceae and Maranta- S.R. JENSEN and B.J. NIELSEN
ments, ceae (Zingiberales) and as the chief storage tissue
storage tissue, which may consist of endosperm, in the family Cyanastraceae (Asparagales).
perisperm, and/or chalazosperm (see below), or
which may be lacking, and
the embryo, which represents the young plant
which has become dormant. The Embryo
Plants are built up mainly from organic molecules reduce exploitation by the less specialized herbi-
The seed coat is here divided into two layers (fol-
and water. In the course of primary metabolism vores.
lowing CORNER, 1976): The embryo consists of the following parts: radicle, they synthesize large amounts of sugars, amino Chemistry has always been used in the classifica-
the testa, which is the outer layer, formed by the hypocotyl, cotyledon and plumule. The radicle is acids, fats and lignans. These are the basic units
outer integument, and tion of plants. The colour of flowers, as well as
the rudimentary root, which in monocotyledons from which are built up the more complex sub-
the tegmen, which is the inner layer, formed by the taste and smell of fruits, seeds and other parts,
ceases to grow at a relatively early stage of devel- stances of the cells such as cellulose, starch and
the inner integument. opment. The hypocotyl is the region of the axis are examples of chemical characters. Until recently
enzymes (protein), end-products that constitute the occurrence of specific compounds has been lit-
These layers vary in relative thickness and in the between the radicle and the cotyledon. The cotyle- most of the organic material in roots, stems, tle known and even our present knowledge is
shape of their cells from one group of monocotyle- don is the first leaf and is often tubular; the exact leaves, flowers and fruits. Most of these basic sub-
dons to another. The epidermis of the testa may relationship of the so-called cotyledon in monocot- based on studies of a very small proportion of
stances are of no taxonomic interest since they existing species.
be black owing to the presence of a characteristic yledons to the two cotyledons in the dicotyledons are ubiquitous in higher plants. Chemists are generally most interested in economi-
carbon-rich compound, phytomelan, which is has not been satisfactorily determined (see p. 44). A certain portion of the compounds formed by cally important plants, and systematic surveys of
known only in taxa of Asparagales (see Fig. 90 M- The plumule is the stem bud and is found at the the primary metabolism is diverted into secondary
P). Rarely, the testa is fleshy, forming a sarcotesta, substantial groups of related species have only re-
base of or inside the cotyledon and emerges from metabolites, giving rise mainly to products in- cently been undertaken. Chemical methods have
as in certain Araceae. it through a lateral pore. It bears the first leaves volved in the basic vital processes of the cell by been improved tremendously in the last three de-
The storage tissue is generally made up chiefly of after the cotyledon and in most monocotyledons acting as regulators (hormones) of the primary me- cades, facilitating the isolation of compounds and
the endosperm, which is surrounded by a thin rem- has an initial distichous phyllotaxy. tabolism. Some, however, may merely accumulate, the determination of their chemical structure. Thus
nant of nucellar tissue. In most orchids endosperm In groups which lack endosperm the Aypocotyl particularly in certain tissues (e.g. the bark). Ex-
is not formed, while in all Alismatiflorae and at a branch of biology that can be called chemotaxon-
may be very thick and fleshy, serving as the storage amples of such compounds are ethereal oils, bitter
least half of the Ariflorae it is absorbed during reservoir. Embryos in this condition are termed omy or biochemical systematics has evolved, with
glycosides and alkaloids. Earlier botanists did not the main purpose of elucidating relationships in
seed development. It is also weakly developed in macropodous and occur in many Alismatiflorae perceive any obvious role for these substances and plant groups, though some studies also consider
the few groups which have a copious (i.e. rich) and Ariflorae. therefore usually regarded them as mere waste
perisperm or chalazosperm. Perisperm is a storage the biological effects of secondary metabolites.
The classification of the different types of embryo products of the primary metabolism. However, it
tissue that consists of nucellar tissue which has un- is outlined in the chapter Distribution of Character The fields of botany, chemistry and pharmacology
has been found that they often make the plants have benefited mutually from this development.
dergone active cell division after fertilization of Conditions (section on Fruit and Seed Charac- poisonous or disagreeable to feeding animals (pre- The Definition of Chemical Characters. For sev-
the ovule. It occurs only in families of Zingiberales ters). dators) and in that way protect them. The presence eral reasons it is not possible to give a precise
of such compounds may thus be of positive selec- definition of a chemical character. If a certain
tive value for the plants and may favour their sur- compound is present in a plant this may be taken
vival. In this context it is usually assumed that to constitute a character condition. But what one
the poisonous or repellent substances are not mere needs to know is whether this compound is present
by-products and that, instead, their production in a whole genus or family under study. Presence
imposes an energy cost on the plant which is “jus- or absence in a single plant, on its own, may be
tified” by the protection gained. Various kinds of taxonomically useless information. Reports of
substances have been shown to be effective against “absence” also have to be viewed in relation to
micro-organisms either by their chemical action the sensitivity of the method of detection used.
(as antibiotics) or by mechanically sealing wounds Absence of a compound in some taxa within a
(gums). Others protect the plant from vertebrate group where the compound is otherwise present
and invertebrate herbivores. may represent a loss of the ability to produce it
There are also cases where herbivores have devel- rather than an ancestral lack of it. Only the verified
oped means of evading the harmful effects of cer- presence of a compound constitutes a character con-
tain compounds, sometimes even becoming depen- dition that is of use for our purpose.
dent on them, so that they seek out the plants The next problem is the significance of the pres-
containing them. Thus the presence of second- ence of a compound (or group of biogenetically
ary compounds is not unambiguously advanta- related compounds). Where, for example, two par-
geous to plants, even though it may effectively ticular families between which there are other con-