The plants in this family have a _curved ovule_, and most frequently
a _kidney-shaped seed_ (generally provided with fine, cuticular,
projecting warts, Fig. 362 _B_), with a _curved, peripheral embryo
enclosing the endosperm which is most frequently floury_ (Figs. 362
_C_, 365 _H_; for exceptions, see Fig. 366); the seeds in all cases
are borne on a _centrally-placed_, and in most cases _free_, placenta
(they are “basal” when there is only 1 ovule in the ovary, Fig. 364).
The flower is regular, hypogynous or perigynous (Fig. 364) (only
rarely epigynous) and usually 5-_merous_. The flower which is most
complete has 5 whorls (S5, P5, A5+5, G2-3–5), as in some genera of
the Caryophyllaceæ (Figs. 360, 361); but from this type it becomes
reduced, the petals and stamens being suppressed, so that finally
5 perianth-leaves, 5 stamens (opposite the perianth-leaves), and 2
carpels (Fig. 361 _F_) only are present; for example, in certain genera
of the _Caryophyllaceæ_, in the _Chenopodiaceæ_, _Amarantaceæ_, and
others. When the number of stamens is increased to more than 5 in the
whorl, it is always possible to show that some of the stamens have been
divided. The number of the carpels and ovules also becomes reduced; in
the highest there is a central placenta, not free in its early stages,
with a large number of ovules; in those which are most reduced there is
only a single ovule, which is placed centrally at the base of the ovary
[Fig. 364]. Somewhat corresponding changes are found in the fruit,
which is a many-seeded _capsule_ in those which have many ovules, but
a one-seeded _nut_ where there is one ovule. In the most reduced forms
the flowers are generally unisexual.--Similar features are also present
in the vegetative parts. Almost all the species are herbaceous, the
leaves are simple and most frequently without stipules. ~The structure
of the stem, especially in Chenopodiaceæ, Amarantaceæ, Nyctaginiaceæ
and others, often differs from that of the ordinary Dicotyledon. In the
woody portion of the stem and root several rings are sometimes formed
which resemble annual rings but which are formed by new cambium-rings
arising outside the old ones which then cease to divide.~
Order 1. =Caryophyllaceæ.= Herbaceous plants, with round, nodose
stem; leaves _opposite_, slightly amplexicaul, simple, with sessile,
undivided, entire lamina; stipules nearly always absent; the
inflorescences are _dichasia_ passing over into unipared scorpioid
cymes. The flowers are regular, ☿ or unisexual, hypogynous or
perigynous, 5-(or 4-) merous with 2–3–4–5 carpels; calyx persistent;
corolla polypetalous. The ovary is unilocular (or originally, and
sometimes also in the later stages, plurilocular below, _e.g._
_Viscaria_), with _free styles_ and 1–several curved ovules on a
_central_, free placenta. The fruit is a nut or a capsule opening
apically with long or short valves (teeth, Fig. 362), equal to or
double the carpels. For the seeds refer to the family. ~In _Dianthus_
the embryo is straight.~
The flowers which are most complete have _Sn_, _Pn_, _An_ + _n_
(obdiplostemonous), _Gn_ where _n_ = 5 (Figs. 360, 361 _A_) or
4 (Fig. 361 _B_); the carpels may be placed opposite to the
sepals (Fig. 360) or opposite to the petals (Fig. 361 _A_, _B_).
Without any change taking place in the position of the other
whorls, the carpels are next found reduced to 2–3–4 (see the
genera); their number may easily be recognised by that of the
styles. This is the construction in the majority of the genera
in the two first groups. _Stellaria media_ differs considerably.
It may have (_a_) the flower as described above, with _G3_;
(_b_), the corolla only absent, or (_c_) only the petal-stamens
(A5 + 0, Fig. 361 _C_), or (_d_) all these as well as some of
the sepal-stamens. The same applies to _Sagina_, _Alsine_,
_Cerastium_, and others, and, finally, a series of genera are
formed, with certain conditions of reduction which have become
constant, and by a gradual series of steps lead to the most
reduced form, which has only 5 sepals and 5 (or even as far as
only 1) sepal-stamens (Fig. 361 _D_, _E_, _F_).--The PETALS in
the _Alsineæ_ are often deeply bifid. The sepal-stamens are most
frequently the longest, and bear nectaries at the base (Fig. 363
_st_). In the most complete forms the ovary has partition-walls
in the lower portion (Fig. 360); these do not, however, reach
to the top, and generally soon disappear. The ovules, when
numerous, are situated on the placenta in as many double rows as
there are carpels. In the number of ovules a reduction from many
to 1 takes place (Fig. 361). A comparison proves that the “free,
centrally placed” placenta is formed by the ventral portion of
the carpels. The single basal ovule in _Herniaria_ (Fig. 364),
_Scleranthus_, and others, is also borne on the carpels.
The vegetative _branching_ is characteristic. One of the leaves
in a pair is formed before the other, and has a more vigorous
axillary bud; these stronger leaves stand in a ¼-spiral, the
fifth above the first one, and the branches are consequently
arranged in the same manner. In the inflorescence, however, it
is the upper or second bracteole (β) whose axillary bud (_w_ in
Fig. 361) is most advanced. The bud of the first bracteole (α)
becomes sometimes entirely suppressed, or in some this bracteole
itself is suppressed.
[Illustration: FIG. 360.--Diagram of _Lychnis_: α, β bracteoles.]
[Illustration: FIG. 361.--_A-F_ Diagrams of flowers of the
Caryophyllaceæ: _A Agrostemma_; _B Sagina_; _C Stellaria_;
_D Corrigiola_; _E Paronychia_; _F Herniaria_.]
The most original type appears to be represented by the Alsineæ.
From this form on one side the Sileneæ, adapted in a higher
degree for insect-pollination, are developed, and on the other
side the Paronychieæ, with various reductions.
=1.= ALSINEÆ, STITCHWORT GROUP. Sepals free, and connected with them
stellately expanded, slightly unguiculate (white or inodorous) petals;
these, however, often become suppressed (Fig. 363). The fruit is a
capsule.
[Illustration: FIG. 362.--_Cerastium arvense_: _A_ fruit; _B_
seed; _C_ section of seed.]
=a.= As many carpels as sepals (4 or 5). _Cerastium_ (Chickweed). The
petals are bifid. Capsule cylindrical, frequently _curved_ at the top,
and opening by 10 teeth (Fig. 362).--~_Malachium_ differs only in the
5-toothed capsule with bifid teeth.~--_Spergula_ (Spurry). The petals
are not bifid, capsule 5-valved; seeds winged. The leaves are linear,
and appear as if placed in large numbers in a whorl, a branch being
situated in the axil of each with leaves placed very close together at
its base; _stipules membranous_.--~Sagina has Sn, Pn, An + n, or An,
Gn, where n = 4 or 5. The corolla is often wanting.~
=b.= 3 (rarely 2) carpels (Fig. 361 _C_). _Stellaria_ (Stitchwort)
has deeply cleft petals. The number of stamens varies (see
above).--_Arenaria_ has entire petals. ~(To this group belong _Alsine_,
_Moehringia_, _Halianthus_, or _Honckenya_ (Fig. 363), which differ
from each other, especially in the form of the seed and number of
the capsular valves.) _Spergularia_ has membranous stipules, as in
_Spergula_.--_Holosteum._~
[Illustration: FIG. 363.--_Arenaria_ (_Halianthus_) _peploides_:
♀-(_A_) and ♂-flower (_B_, _C_).]
=2.= PARONYCHIEÆ (Figs. 361 _D_, _E_, _F_; 364). Small, greenish
plants. The leaves, in the majority, are opposite, with _membranous
stipules_. The flowers are most frequently arranged in small
_dichasia_; they are small and insignificant, perigynous (Fig. 364)
or hypogynous. The corolla is in most cases wanting, and when present
is very small; in general the calyx-stamens are developed, but the
corolla-stamens may be represented by small scales (Fig. 364). Ovary
most frequently with 1 ovule. Fruit, a _nut_, rarely a capsule; it is
enclosed by the strongly perigynous floral axis (torus).
_Scleranthus_ (Knapwell) is perigynous with bell-shaped torus; no
corolla; corolla-stamens are wanting or rudimentary; some calyx-stamens
may also be absent.--_Corrigiola_ (Fig. 361 _D_); _Illecebrum_;
_Paronychia_ (Fig. 361 _E_); _Herniaria_ (Figs. 361 _F_, 364).
=3.= SILENEÆ, PINK OR CARNATION GROUP. This has a _gamosepalous_ calyx
and unguiculate, white or red, petals, with _outgrowths_ (_ligule_,
_corona_, _paracorolla_) at the throat of the corolla. These structures
are not found in the other groups, and are merely outgrowths at
the junction of the limb and claw. The corolla, stamens and ovary
are frequently raised above the calyx, upon a lengthened internode
(_gynophore_). The flower has S5, P5, A5 + 5; fruit a capsule with many
seeds.
=a.= 5-(rarely 3–4) carpellate ovary.--_Lychnis_ (Campion, Fig. 360).
The corolla is longer than the calyx; corona present. The capsule
is 10- or 5-toothed, completely 1-chambered or 5-chambered at the
base,--the genus has been divided accordingly into several genera:
_Melandrium_, _Lychnis_, _Viscaria_. ~Some species are unisexual by
the abortion of stamens or carpels (_L. vespertina_, _diurna_).~
_Agrostemma_ (_A. githago_, Corn-cockle, Fig. 361 _A_) has a
long-toothed calyx, the teeth exceeding the corolla; corona absent;
5-toothed capsule.
[Illustration: FIG. 364.--_Herniaria glabra_: _a^1_ flower; _b^1_
longitudinal section through the flower; _c^1_ stigma with two
pollen-grains.]
=b.= Tricarpellate.--_Silene_ (Catch-fly). Six-toothed capsule; corona
present in the majority.--~_Cucubalus_ has berry-like fruits which
finally become dry but do not dehisce.~
=c.= Bicarpellate (2 styles, 4-toothed capsule).--_Dianthus_
(Pink); at the base of the calyx 1–several pairs of floral-leaves
are situated; corona absent. The _straight embryo_ is a peculiar
exception.--_Gypsophila_ has a campanulate, open calyx, 5-nerved,
membranous between the nerves; corona absent; the flowers are generally
small and numerous, in a large, paniculate dichasia.--_Saponaria_
(Soapwort) has corona.
POLLINATION. _Alsineæ_ has ordinary nectaries at the base of
the calyx-stamens (Fig. 336): they are frequently protandrous
but may often, in the absence of cross-pollination (in the
less conspicuous species) pollinate themselves. Their open
flowers are accessible to many kinds of insects (particularly
flies and bees). _Gynodiœcious_ flowers are found in several
species, and the ☿-flowers are then generally more conspicuous
than the ♀-flowers. That the ♀-flowers have descended from
☿-flowers is seen by the large staminodes found in them (Fig.
363). _Arenaria peploides_ is diœcious (Fig. 363). The _Sileneæ_
are as a rule adapted for pollination by insects with long
probosces--especially butterflies,--and they are frequently
protandrous, so that at first the calyx-stamens open, later
on the corolla-stamens, then the stigmas expand. The honey is
secreted by a ring-like nectary round the base of the ovary or
by nectaries at the base of the stamens. Some only blossom and
emit scent at night or in the evening (_Lychnis vespertina_,
_Silene nutans_, _Saponaria officinalis_) and, like other
night-flowers, are of a white or pale colour.
DISTRIBUTION. 1,100 species, especially in temperate climates,
fewer in the colder zone, less still in the Tropics. The
Paronychieæ are especially found in dry, sandy fields.
USES. “Soap-root” (with _Saponin_, forming a lather in water)
from _Saponaria officinalis_ was formerly officinal, and
_Gypsophila struthium_. The seeds of _Agrostemma githago_ are
said to be poisonous.--The following are ornamental plants:
species of Pinks (_D. caryophyllus_, garden Pink, often
with double flowers; _D. barbatus_, _plumarius_, _etc._).
_Lychnis_, _Gypsophila_, _Silene_, _Cerastium_ (_C. tomentosum_
as edging for borders), _Saponaria officinalis_ (often
coronate).--_Spergula arvensis_ is sometimes cultivated.
Order 2. =Amarantaceæ.= The flowers are essentially the same as
in the _Chenopodiaceæ_ and the extremely reduced Caryophyllaceæ
(Fig. 361 _F_); they are regular, hypogynous, generally ☿,
have 5 free (rarely slightly united) perianth-leaves; in front
of these 5 stamens, which _are often united_ at their base
into a shorter or longer tube and have stipule-like teeth
between them (the division _Gomphreneæ_ has 2-locular anthers,
each of which opens longitudinally); and a 2–3 carpellate
gynœceum with one loculus and most frequently one, more rarely
several ovules; the fruit is a nut, more rarely (in _Celosia_,
_Amarantus_, _Gomphrena_) a capsule, dehiscing irregularly,
or like a pyxidium. The characters which especially separate
them from the allied orders are found in the perianth. The
perianth-leaves are not green and herbaceous, but _membranous,
dry, and often coloured_; they are frequently produced into a
bristle or awn; they have also both subtending floral-leaves
and _2 large bracteoles similar to the perianth_; all these dry
leaves persist without alteration after the withering of the
flower.--The flowers are without scent. They are arranged in
spike- or capitulum-like inflorescences; sometimes placed singly,
sometimes aggregated in the panicle-like inflorescences; in
others, on the contrary, in dichasia. The majority are herbs,
some are shrubs. The leaves are scattered, or opposite, but
always simple and without stipules; some are smooth, others
hairy.
450 species; especially in the Tropics, principally S. Am. and
E. Ind.: few are found outside these countries.--Only a few
are used; some, chiefly E. Indian species, are cultivated as
ornamental plants: _Amaranthus_ (Fox-tail); _Gomphrena globosa_;
_Celosia cristata_ (Cock’s-comb) remarkable for its fasciated
inflorescence; _Alternanthera_. Some are employed as culinary
plants in the Tropics, and in a few of the E. Indian species the
seeds are farinaceous, and used for food.
Order 3. =Chenopodiaceæ.= Generally herbaceous plants like the
Caryophyllaceæ, but the leaves are arranged spirally (except
_Salicornia_), and are simple, exstipulate; they are generally fleshy
and like the stem “mealy,” that is, covered with small hairs, whose
large spherical terminal cell readily falls away; otherwise they are
seldom hairy. The inflorescences are generally flower-clusters borne in
panicles. Bracteoles generally absent. Flowers generally _unisexual_:
with the single exception of _Beta_ the flowers are hypogynous; they
are regular, small and inconspicuous, with _single, green_, 5-leaved,
but _more or less united_ perianth; 5 stamens opposite the perianth,
and a _2–5-carpellate, unilocular_ ovary with 1 basal, curved ovule;
but in some genera the number of the perianth-leaves and stamens is
reduced to 3–2–1–0. The fruit is generally a _nut_,--thus this flower
and fruit are the same as in the reduced Caryophyllaceæ (Fig. 361
_F_). The seed is similar to that generally found in the family (for
exceptions see the genera).
The floral diagram most frequently present is the same as
in Fig. 361 _F_. There is no indication of corolla or of
corolla-stamens, which may be supposed to have belonged
to the plant, but which are now entirely and completely
suppressed. This order appears to have been an offshoot from
the Caryophyllaceæ.--The perianth persists after the withering
of the flower, and envelopes the nut; it is very variable, and,
together with the position of the seed, the form of the embryo,
the sex of the flowers, etc., gives the characters of the genera.
=1.= CHENOPODIEÆ, GOOSEFOOT GROUP (Fig. 365), has ☿ (or polygamous)
flowers, with regular 5-parted perianth (_C_); the embryo is ring-like
(_H_). The leaves have the ordinary flat forms.--_Chenopodium_
(Goosefoot). The flower is hypogynous, and the fruit (which is
compressed) perfectly free; Mulberry-like collections of fruits are
formed in some species (sub-genus _Blitum_) by the perianth becoming
finally fleshy and coloured.--_Beta_ (Beet, Mangold, Fig. 365) differs
from all genera in the perianth, which finally becomes cartilaginous,
being epigynous (_D_). Small, most frequently 2–3-flowered clusters
without bracteoles, situated in a long, interrupted axis (_A_, _B_);
the flowers and fruits in each cluster are more or less united
individually, and fall off together--they are commonly known as seeds
(_E_, _F_). The seed lies horizontally.--_Hablitzia_ (_H. tamnoides_).
[Illustration: FIG. 365.--_Beta vulgaris._]
[Illustration: FIG. 366.--_Salsola soda_: embryo.]
=2.= SALSOLEÆ, SALTWORT GROUP, has cylindrical or semi-cylindrical
leaves. Perianth as in the preceding group; the fruit is most
frequently compressed. The two first mentioned genera differ from
most of the others in the order in having a spirally-coiled, and not
a ring-like embryo, so that the endosperm is slight or wanting (Fig.
366). These plants are sometimes placed as a group by themselves,
SPIROLOBEÆ--in contradistinction to which the others are termed
CYCLOLOBEÆ.--_Salsola_ (Saltwort); leaves subulate, with spiny tips;
the flowers have 2 spinous bracteoles: during the ripening of the
fruit a tough leathery wing is developed transversely to the back of
the perianth.--~_Chenopodina_ deviates from _Chenopodium_ chiefly in
the embryo and want of endosperm.--_Kochia_ has a somewhat similar
perianth to _Salsola_, but a ring-like embryo; it differs from the
others in being hairy.~
=3.= SALICORNIEÆ, GLASSWORT GROUP. _Salicornia_ (Glasswort) has a very
different appearance. The stems are succulent, jointed, and almost
leafless; the leaves opposite, very small, sheath-like and connate;
there is a depression in the axil of each leaf, in which a small
3-flowered dichasium without bracteoles is sunk; the flowers have
a trimerous perianth, 1–2 stamens and 1 carpel. No endosperm. _S.
herbacea_ on clayey beaches.
=4.= ATRIPLICEÆ. This group has most frequently unisexual flowers;
the ♂-flower has a 4–5 partite perianth, but the ♀-flower differs
from it. _Atriplex_ is monœcious or polygamous, the ♀-flower is
naked, but has 2 large, herbaceous bracteoles which expand during the
ripening of the fruit, and often become warted and fringed, enveloping
the _compressed nut_. ~The section _Dichospermum_ has two kinds of
♀-flowers, one like those just described, the other similar to the
_Chenopodium_-flowers, which have been deprived of their stamens, and
the fruits of which are _depressed_, not pressed together from the
sides; some (_e.g. A. hortensis_) have even three kinds of nuts.
All the flowers of _Atriplex_, which present vertical fruits, are
accessory shoots, which stand beneath the ordinary flower-clusters, a
rather singular relation.~--_Spinacia_ (Spinach) is diœcious; ♂-flower:
perianth, 4 (-5); stamens, 4 (-5); ♀-flower: tubular, 2–4-partite
perianth, hardening during the ripening of the fruit, and uniting with
the compressed nut; in _S. oleracea_, it also forms _thorns_; 4 long
stigmas.--_Halimus_ has the 2 long bracteoles almost entirely united
and ultimately adhering firmly to the fruit.
=5.= BASELLEÆ. A somewhat exceptional group with more
or less perigynous flowers and 2 bracteoles. _Basella_,
_Boussingaultia_, _Ullucus_. The perianth is sepaloid; ovary
1-ovuled. In _Basella_ the perianth is fleshy, enveloping the
nut, and the cotyledons are so rolled together that a tranverse
cut divides them in two places (as in Spirolobeæ). Herbaceous
climbing plants.
POLLINATION. Wind-and self-pollination, as far as is known;
the insignificant flowers, devoid of honey, appear to exclude
insect-pollination.--520 species. Most of them are annual
(out of 26 native species only 5 are perennial); inhabiting
salt-marshes and salt-steppes, and growing as weeds (most
frequently on garden or field soil containing manure) in this
country, especially species of _Chenopodium_ and _Atriplex_.
The majority are found outside the Tropics, and play a very
important part, for example, in the Asiatic salt-steppes. They
grow gregariously in large masses.
USES. Comparatively few. The only important one is _Beta
vulgaris_ (from the Mediterranean basin), with its different
varieties, viz. Beet-root, Cattle-beet or Red-beet, Sugar-beet,
and others. These are biennial, making in the first year a root
which acts as a reservoir of reserve material, with a rosette
of leaves, and in the second year using this material in the
production of a long stem, leaves and flowers. The primary
root has been developed by cultivation into a very thick and
fleshy tap-root; its mode of increase in thickness deviates
from that of other roots, concentric rings of vascular bundles
being formed from a cambial ring developed outside the previous
ring. In this way several rings of vascular bundles separated
by medullary rays, alternating with rings of parenchyma, may
be found in the root of a Beet. Besides _Beta vulgaris_, var.
_hortensis_ (Beet-root), the following are also cultivated: var.
_cicla_ (Leaf-beet, “Mangold,” or “Roman Spinach ”), _Spinacia
oleracea_ and _Atriplex hortensis_ as Spinach; a form of the
latter and of Spinach are grown as ornamental plants. The tubers
of _Ullucus tuberosus_ are used as potatoes; _Chenopodium
quinoa_, in Chili and Peru, is an important farinaceous plant.
Soda is made from some (_Salsola kali_, _Chenopodina maritima_
and others). Aromatic properties are rare: _Chenopodium
ambrosioides_ and _botrys_.
Order 4. =Batidaceæ.= _Batis maritima_, a bushy West Indian
maritime plant.
Order 5. =Phytolaccaceæ.= The ☿ (sometimes unisexual), regular,
sometimes slightly perigynous flowers are inconspicuous
and have a single sepaloid or coloured 4–5-leaved perianth
(generally united at the base); stamens either in 1 whorl in the
spaces between the perianth-leaves or in 1 whorl opposite the
perianth-leaves, or in 2, one of which alternates with these;
but the number may be increased by the splitting of one or of
both the whorls to as many as 10–15–20–25. _Carpels_ sometimes
only one, sometimes _many_ (4–10) placed in a whorl, either free
or united into a gynœceum with a corresponding number of loculi
in the ovary; but in all cases _each carpel bears only its own
style and 1 ovule_. The fruit is a _berry_ (or nut, capsule, or
schizocarp).--Mostly herbs or herbaceous shrubs, with scattered,
simple leaves without stipules (_Petiverieæ_; have stipules).
Inflorescences, most frequently _racemes_ or spikes, which in
some instances are apparently placed opposite to a leaf, being
displaced by a more vigorous growth of the axillary bud. Embryo
always bent.--_Petiveria_ has a straight embryo with rolled
cotyledons.--_Phytolacca_, _Pircunia_, _Microtea_, _Seguieria_,
_Rivina_ (Pr4, A4, G1; berry), and others.
The following plant is, with some doubt, placed near this
order: _Thelygonum cynocrambe_; monœcious. ♂-flowers: perianth,
2-leaved; stamens indefinite. ♀-flowers: perianth-leaves united,
3-toothed; G1, style gynobasic. Fruit a drupe. An annual plant;
Mediterranean. Branching anomalous.
About 90 species; in tropical and temperate countries,
principally America and Africa.--The red juice in the fruits,
especially of _Phytol. decandra_, is used for colouring wine.
Order 6. =Portulacaceæ= (=Portulacas=). The flowers are regular
(except _Montia_), hypogynous (except _Portulaca_) and ☿. The diagram
which applies to the majority of genera is that in Fig. 367, but with
all the 5 stamens completely developed: it may be considered as the
Chenopodiaceous diagram with the addition of 2 _bracteoles_ in the
median line (_m-n_, these by some are considered as sepals), and with
a petaloid perianth (usually designated “corolla”). The “petals” fall
off very quickly, and are sometimes wanting. Most frequently 5 stamens,
situated opposite the “petals,” but in other genera the number varies;
_Montia_ has only 3 stamens (by suppression of the two anterior and
lateral, Fig. 367), others again have more than 5, some a large and
indefinite number. This may be explained partly by the appearance of
a second whorl of stamens alternating with the first, and partly by
the splitting (dédoublement) of the stamens. Gynœceum most frequently
tricarpellate, ovary unilocular with 1–several basal ovules (sometimes
on a branched placenta, as in certain _Caryophyllaceæ_). The fruit is
a _capsule_, more rarely a nut.--The majority are annual herbaceous
plants with scattered, entire leaves, often fleshy and smooth, with or
without rudimentary stipules (dry, membranous, modified into hairs).
Inflorescence cymose.
[Illustration: FIG. 367.--_Montia._