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Leptestheria,[29] G. O. Sars—Carapace compressed, oblong.
Rostrum with a movable spine; thoracic limbs with
accessory lappet on the exopodite. L. siliqua, G. O. Sars—
Cape Town.
Cyclestheria,[30] G. O. Sars. C. hislopi, Baird—Queensland,
India, East Africa, Brazil.
Sub-Order 2. Cladocera.
The Cladocera are short-bodied Branchiopods, with not more than six
pairs of thoracic limbs. The second antennae are important organs of
locomotion, and are nearly always biramous; the first antennae are small,
at least in the female; the second maxillae are absent in the adult. The
carapace may extend backwards so as to enclose the whole post-cephalic
portion of the body, or may be reduced to a small dorsal brood-pouch,
leaving the body uncovered.
The Cladocera or “Water-fleas” are never of great size; Leptodora
hyalina, the largest, is only about 15 mm. long, while many Lynceidae are
not more than 0·1 or 0·2 mm. in length.
The head is bent downwards in all the Cladocera, so that parts which
are morphologically anterior, such as the median eye and the first
antennae, lie ventral to or even behind the compound eyes and the second
antennae (cf. Fig. 10).
The compound lateral eyes fuse at an early period of embryonic life, so
that they form a single median mass in the adult, over which a fold of
ectoderm grows, to make a chamber over the eye, like that found in the
Limnadiidae, except that it is completely closed. The fused eyes are
generally large and conspicuous; in some deep-water forms the retinular
elements of the dorsal portion are larger than those of the ventral (e.g.
Bythotrephes, Fig. 13). In one or two species which live at very great
depths, or in caves, the eyes are altogether absent.
The appendages of the head are fairly uniform, the most variable being
the first antennae. In the females of many genera the first antennae are
short and immovable, consisting of a single joint, with a terminal bunch
of sensory hairs, and often a long lateral hair, as in Simocephalus (Figs. 9,
10), Daphnia, etc. In the female Moina (Fig. 16) they are movable, as they
are in Ceriodaphnia and some others; in Bosmina (Fig. 22) and many
Lyncodaphniidae they are elongated and imperfectly divided into joints
by rings of spines, while in Macrothrix they are flattened plates. In the
males the first antennae are elongated and mobile (cf. Figs. 11, 19).
The second antennae, the chief
organs of locomotion, are biramous
in all genera except Holopedium;
the number of joints in each ramus,
and the number of the long plumose
hairs with which they are provided,
are remarkably constant in whole
series of genera, and are therefore
useful for purposes of classification.
The creatures row themselves by
quick strokes of these appendages,
the movement being slow and
irregular in the rounder forms, such
as Simocephalus or Daphnia, rapid
and well directed in such elongated
Fig. 9.—Simocephalus vetulus, female.
Ventral view, without the carapace; A1, lacustrine forms as Bythotrephes or
A2, first and second antennae; For, head; Leptodora.
Md, mandible; Te, telson; I-IV, first to The mandibles have no palp; the
fourth thoracic appendages. first maxillae are very small, and the
second maxillae are absent (Fig. 9).
The carapace varies very much. In most genera (the Calyptomera of
Sars) it is a large, backwardly-projecting fold of skin, bent downwards at
the sides so as to form a bivalve shell, enclosing the whole post-cephalic
portion of the body, as in Simocephalus (Fig. 10). The eggs are laid into
the space between the carapace and the dorsal part of the thorax, both the
carapace and the thorax itself being often modified for their protection
and nutrition. In a few forms, the Gymnomera of Sars, the carapace
serves only as a brood-pouch, which is distended when eggs are laid, but
collapses to an inconspicuous appendage at the back of the head when it
is empty (e.g. Leptodora, Fig. 24, Bythotrephes, Fig. 13). In the
Calyptomera the surface of the carapace is frequently provided with a
series of ridges, which may be parallel, rarely branching, as in
Simocephalus; or in two sets which cross nearly at right angles, as in
Daphnia; or so arranged as to form a hexagonal pattern, as in
Ceriodaphnia. In a few forms the whole surface is irregularly covered
with spines or scales. The hinder edge of the carapace is often produced
into a median dorsal spine (Daphnia, Fig. 19), or more rarely there are
two spines, one at each ventro-lateral corner (Scapholeberis, Fig. 20).
Fig. 10.—Simocephalus vetulus, × 30. Side view of female, showing the
arrangement of the principal organs. A.2, Second antenna; C.S,
cervical suture; E, fused compound eyes; H, heart; L, forwardly-
directed gastric caeca; N, dorsal organ.
The cuticle of the carapace is often separated from that of the head by a
cervical suture, as in Simocephalus (Fig. 10, C.S.) and near the line of
demarcation many forms exhibit patches of glandular ectoderm which
seem to be homologous with the dorsal adhesive organs of the
Limnadiidae. The commonest condition is that of a median dorsal pit
(Fig. 10, N.) by means of which the animal can fix itself to foreign objects.
Certain forms may remain for long periods of time attached by the dorsal
organ to plants, or to the sides of an aquarium, the only movement being
a slow vibration of the feet, by which a current of water, sufficiently rapid
for respiratory purposes, is established round it.[31] In Sida crystallina
(Fig. 11) the dorsal organ is represented by three structures; in front there
is a median raised patch (N.m) of columnar ectoderm, containing
concretions like those described in the Branchipodidae, and behind this is
a pair of cup-shaped organs (N.e), with raised margins.
The fold of skin which forms the carapace contains the coils of the
single pair of kidneys, and it forms an important organ of respiration,
partly from the great size of the blood-vessels it contains, and partly from
the presence of red, blue, or brown
respiratory pigments in the tissue of
the skin itself.
In most Cladocera the cuticle of
the carapace is cast at every ecdysis,
with that of other parts of the body;
but in Iliocryptus and a few others it
remains after each moult, giving the
carapace an appearance of “lines of
growth,” like that seen in many
Limnadiidae.
The segmentation of the body
behind the head is obscure, but we
can generally recognise (1) a thorax,
of as many segments as there are
pairs of limbs; (2) an abdomen of
three segments; and (3) a telson.
The thoracic limbs of the
Calyptomera are flattened, and
resemble those of the Phyllopoda; as
a type we may examine the third
thoracic limb of Simocephalus (Fig.
12, C), in which the axis bears a
large setose gnathobase (Gn) on its
inner edge, followed by two small
endites; the terminal process, or
exopodite (Ex) is a large flattened
Fig. 11.—Sida crystallina, male, × 27. plate, with six long plumose hairs on
Oxford. A.1, Elongated first antenna; N.e, its edge. The outer margin of the
paired element of dorsal organ; N.m, axis bears a bract (Br) and an
median element of dorsal organ; Te, epipodite.
testes; ♂, opening of vas deferens.
In Simocephalus, as in the other
Daphniidae, there are five pairs of
thoracic limbs, of which the third and fourth are alike; in the female each
limb of the first pair consists of a jointed axis, with strong biting hairs on
the inner border, and a rudimentary epipodite (Fig. 12, A), the second
limb being more like the third, but with a more prominent gnathobase
and a narrower exopodite (B), while the limbs of the fifth pair have the
gnathobase and the exopodite filamentous (D).
In the Sididae there are six pairs of thoracic limbs, which are nearly
alike in the female; in the Bosminidae there are six pairs, the first two
modified for prehension, the last much reduced.
The summer-eggs are always carried until they are hatched by the
parthenogenetic female which produces them. The brood-pouch is the
space between the dorsal wall of the thorax and the carapace. This space
is always more or less perfectly
closed at the sides by the pressure of
the carapace against the body, and
behind by vascular processes from
the abdominal segments (Figs. 10,
16, etc.). The presence of a large
blood-sinus beneath the dorsal wall
of the thorax and in the middle line
of the carapace suggests the
possibility that some special
nutrient substances may pass from
the body of the parent into the Fig. 17.—Moina rectirostris, ♀ , × 40,
brood-chamber, and in some showing the ephippial thickening of the
species the thoracic ectoderm is carapace which precedes the laying of a
specially modified as a placenta. In winter-egg.
Moina (Fig. 16) the dorsal wall of
the thorax is produced into a dome,
covered by a columnar ectoderm, which contains a dilatation of the dorsal
blood-sinus; and in this form it has been shown that the fluid in the
brood-pouch contains dissolved proteids. Associated with the apparatus
for supplying the brood-pouch with nutriment is a special apparatus for
closing it, in the form of a raised ridge, which projects from the back and
sides of the thorax and fits into a groove of the carapace.
A somewhat similar nutrient apparatus exists in the Polyphemidae,
where the edges of the small carapace are fused with the thorax, so that
the brood-pouch is completely closed, and the young can only escape
when the parent casts her cuticle. In some genera of this family (e.g.
Evadne) the young remain in the parental brood-pouch until they are
themselves mature, so that when they are set free they may already bear
parthenogenetic embryos in their own brood-pouches.
The winter-eggs are fertilised in
the same part of the carapace of the
female in which the parthenogenetic
eggs develop, but after fertilisation
they are thrown off from the body of
Fig. 18.—Newly-cast ephippium of the mother, either with or without a
Daphnia, containing two winter-eggs. protective envelope formed from the
cuticle of the carapace. The eggs of
Sida are surrounded by a thin layer
of a sticky substance, and when cast out of the maternal carapace they
adhere to foreign objects, such as water-weeds; those of Polyphemus have
a thick, gelatinous coat; in Leptodora and Bythotrephes the egg secretes a
two-layered chitinous shell. In these forms the cuticle of the parent is not
used as a protection for the winter-eggs, although it is generally, if not
invariably, thrown off when the eggs are laid. In the Lynceidae the cuticle
is moulted in such a way that the winter-eggs remain within it, at least for
a time; the cuticle is occasionally modified before it is thrown off; thus in
Camptocercus macrurus the cuticle of the carapace, in the region of the
brood-pouch, becomes thickened and darkly coloured, forming a fairly
strong case round the eggs. The modification of the cuticle round the
brood-pouch is much more pronounced in the Daphniidae, where it leads
to the formation of a saddle-shaped cuticular box, the “ephippium,” in
which the winter-eggs are enclosed. The ripening of a winter-egg in the
ovary of a Daphnia is accompanied by a great thickening of the cuticle of
the carapace (cf. Fig. 18), so that a strong case is formed in the position of
the brood-pouch. The winter-eggs are laid between the two valves of this
case, and shortly afterwards the parent moults. The eggs are retained
within the ephippium, from which the rest of the cuticle breaks away (Fig.
18). After separation, the ephippium, which contains a single egg (Moina
rectirostris) or usually two (Daphnia, etc.), either sinks to the bottom, as
in Moina, or floats.
The winter-eggs usually go through the early stages of segmentation
within a short time after they are laid, but after this a longer or shorter
period of quiescence occurs, during which the eggs may be dried or frozen
without injury. The sides and floor of a dried-up pond are often crowded
with ephippia, containing winter-eggs which develop quickly when
replaced in water; and the resting-stage of winter-eggs produced in
aquaria can often be materially shortened by drying the ephippia which
contain them, though such desiccation does not appear to be necessary
for development. Under normal conditions large numbers of winter-eggs
remain quiescent through the winter and hatch in the following spring.
The individual developed from a sexually fertilised winter-egg is
invariably a parthenogenetic female: the characters of the succeeding
generations differ in different cases.
In a few forms, of which Moina is the best known, the parthenogenetic
female, produced from a winter-egg, may give rise to males, to sexual
females, and to parthenogenetic females, so that the cycle of forms which
intervene between one winter-egg and the next is short. A sexual female
produces one or two winter-eggs, and if these are fertilised they are
enclosed in an ephippium and cast off; if, however, the eggs when ripe are
not fertilised, they atrophy, and the female produces parthenogenetic
eggs, being thenceforward incapable of forming sexual “winter” eggs. An
accidental absence of males may thus lead to the occurrence of
parthenogenesis in the whole of the second generation. The regular
production of sexual individuals in the second generation from the
winter-egg appears to depend on a variety of circumstances not yet
understood. Mr. G. H. Grosvenor tells me that Moina from the
neighbourhood of Oxford may give rise to several successive generations
of parthenogenetic individuals, when grown in small aquaria.
In the greater number of Daphniidae, the parthenogenetic female,
produced from a winter-egg, gives rise only to parthenogenetic forms, and
it is not until after half a dozen parthenogenetic generations have been
produced that a few sexual forms appear, mixed with the others. Such
sexual forms are fairly common in April or May in this country; they
produce “winter” eggs and then die, the generations which succeed them
through the summer being entirely parthenogenetic. In late autumn
sexual individuals are again produced, giving rise to a plentiful crop of
winter-eggs, but many parthenogenetic females are still found, and some
of these appear to live and to reproduce through the winter.
In Sida, in the Polyphemidae and Leptodoridae, and in most of the
Lynceidae, sexual individuals are produced only once in every year, while
in a few forms which inhabit great lakes the sexual condition occurs so
rarely that it is still unknown.
Weismann[33] has pointed out that the sexual forms, with their property
of producing eggs which can endure desiccation, recur most frequently in
species such as Moina, which inhabit small pools liable to be dried up at
frequent intervals, while the species which produce sexual forms only
once a year are all inhabitants either of great lakes which are never dry, or
of the sea. Many suggestions have been made as to the environmental
stimulus which induces the production of sexual individuals, but nothing
is definitely known upon the subject.
We have said that even in those generations which contain sexual males
and females there are always some parthenogenetic individuals; there is
therefore nothing in the behaviour of Daphniidae, either under natural
conditions or when observed in aquaria, to suggest that there is any
natural or necessary limit to the number of generations which may be
parthenogenetically produced.
The parthenogenetic Daphniidae are extremely sensitive to changes in
their surroundings; small variations in the character and amount of
substances dissolved in the water are often followed by changes in the
length of the posterior spine, in the shape and size of crests on the head,
and in other characters affecting the appearance of the creatures, so that
the determination of species is often a matter of great difficulty. It is
remarkable that the green light which has passed through the leaves of
water-plants appears to have a prejudicial effect upon some species.
Warren has shown that Daphnia magna reproduces more slowly when
exposed to green light, and that individuals grown in this way are more
readily susceptible to injury from the presence of small quantities of salt
(sodium chloride) in the water than individuals which have been exposed
to white light.
The majority of the Cladocera belong to the floating fauna of the fresh
waters and seas; a few are littoral in their habits, clinging to water-weeds
near the shore, a very few live near the bottom at considerable depths, but
the majority belong to that floating fauna to which Haeckel gave the name
of “plankton.” The Crustacea are an important element in the plankton,
whether in fresh waters or in the sea, the two great groups which
contribute most largely to it being the Cladocera and the Copepoda. For
this reason it will be more convenient to discuss the habits and
distribution of individual Cladocera and Copepoda together in a chapter
specially devoted to the characters of pelagic faunas (cf. Chap. VII.). We
will only add to the present chapter a table of the families with a diagnosis
of the British genera.
ii. First antennae of female long, mobile. Moina, Baird (Figs. 16, 17,
21): median eye absent. Posterior margin of carapace without a
spine.
Fam. 4. Bosminidae: feet
equidistant, five or six pairs; the
first antennae of the female
immobile, with sense-hairs
arranged in rings, not forming an
apical tuft. The intestine uncoiled;
no caeca. Bosmina, Baird (Fig. 22).
Fam. 5. Lyncodaphniidae: four,
five, or six pairs of equidistant
thoracic limbs; the first two pairs
prehensile. First antennae of female
Fig. 22.—Bosmina sp., female, × about mobile, with apical sense-hairs.
80. Lake Constance. Intestine coiled or straight.
i. Four pairs of thoracic limbs.
Lathonura, Lilljeborg.
ii. Five pairs of thoracic limbs.
a. The four-jointed ramus of the second antenna with four
swimming hairs. Macrothrix, Baird: the first antennae of the
female flattened, curved. The intestine simple, straight.
Streblocerus, Sars: first antennae of the female very little
flattened, curved
backwards and outwards.
The intestine coiled, the
stomach with two
forwardly-directed caeca.
b. The four-jointed ramus
of the second antenna
with only three swimming
hairs. Drepanothrix, Sars.
iii. Six pairs of thoracic limbs;
the labrum provided with
an appendage.
Acantholeberis, Lilljeborg:
appendage of labrum long,
pointed, and setose.
Intestine without caecum.
Ilyocryptus, Sars:
appendage of the labrum
short, truncated. Intestine Fig. 23.—Acroperus leucocephalus, ×
with a caecum. about 35. Oxford.