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Family 1. Syllidae see p. 306 Family 8. Amphinomidae see p. 318
" 2. Hesionidae " 308 " 9. Eunicidae " 318
" 3. Aphroditidae " 309 " 10. Glyceridae " 320
" 4. Phyllodocidae " 313 " 11. Sphaerodoridae " 320
" 5. Tomopteridae " 315 " 12. Ariciidae " 321
" 6. Nereidae " 315 " 13. Typhloscolecidae " 321
" 7. Nephthydidae " 317
Sub-Order 2. Spioniformia.
Family 1. Spionidae see p. 321 Family 4. Magelonidae see p. 325
" 2. Polydoridae " 323 " 5. Ammocharidae " 325
" 3. Chaetopteridae " 323
Sub-Order 3. Terebelliformia.
Family 1. Cirratulidae see p. 325 Family 3. Ampharetidae see p. 330
" 2. Terebellidae " 327 " 4. Amphictenidae " 330
Sub-Order 4. Capitelliformia.
Family. Capitellidae, see p. 331.
Sub-Order 5. Scoleciformia.
Family 1. Opheliidae see p. 331 Family 4. Scalibregmidae see p. 334
" 2. Maldanidae " 332 " 5. Chlorhaemidae " 334
" 3. Arenicolidae " 333 " 6. Sternaspidae " 335
Branch B. Cryptocephala.
Sub-Order 1. Sabelliformia.
Family 1. Sabellidae see p. 336 Family 3. Amphicorinidae see p. 339
" 2. Eriographidae " 338 " 4. Serpulidae " 339
Sub-Order 2. Hermelliformia.
Family. Hermellidae, see p. 341.
In Aphroditidae and certain Amphinomidae the body is more or less oval in shape.
In Lipobranchius and Sternaspis it is grub-like, short, and cylindrical, with rounded
ends; in the former it is difficult to distinguish head and tail, or dorsal and ventral
surfaces.
The segments composing the trunk may be all alike, or may constitute two more or
less sharply marked regions, the thorax and abdomen, differing in the character of
the chaetae, or in their arrangement, or in some other way, as in the Sabelliformia
and the Capitelliformia.
The posterior extremity is generally more or less narrowed, and most of the
Nereidiformia are provided with special elongated cirri, borne by the anal segment.
In the Maldanidae and others the body terminates in a funnel, at the bottom of
which is placed the anus. Only in a few cases is the anus not terminal; in
Notopygos and other Amphinomidae, as well as in some species of Polynoë, it is
dorsal.[313] In Sabellaria and Pectinaria the hinder end of the body undergoes
great degeneration; in the former it is achaetous, but cylindrical and bent forwards
alongside the body (Fig. 131). In Pectinaria (Fig. 177), this region, which is called
the "scapha," is leaf-like, and serves to close the narrower end of the tube in which
the worm lives. Arenicola marina, and some Terebellids have no chaetae in the
hinder, narrower part of the body.
In the family Serpulidae one (rarely two) of the most dorsally placed gill filaments
is enlarged terminally, and acts as a stopper or "operculum," which closes the
mouth of the tube when the animal withdraws into it. Further, in Spirorbis this
operculum is grooved on one side, and serves as a brood pouch in which the eggs
undergo development (Fig. 184, p. 341). It will be seen, therefore, that the palps
may be very important organs for the life of the worm, and they are no less
interesting to the comparative anatomist, serving as they do as an excellent
illustration of the various uses which Nature finds for one and the same organ.
In the other sub-Orders the prostomium carries neither palps nor tentacles.
In the Cryptocephala there is never more than a single pair of tentacles, and these
are generally reduced to a group of sensory cells, though in Sabellaria they retain
a considerable size.
In a few genera, such as Aphrodite, Nephthys, Capitella, the first postoral segment
is distinguished from the succeeding segments only by its position with regard to
the mouth (Fig. 132) and by its smaller size. But in the remainder of the
Polychaeta, with here and there an exception, the peristomium is achaetous in the
adult.[319]
In a few cases, such as the Chlorhaemids and Sternaspis, and to a slight degree
in Arenicola, the "head" and even the anterior part of the worm is capable of being
withdrawn into the body.
In most Annelids the chaetae are in two bundles on each side, but there are
certain families in which the dorsal bundle, and even the notopodium itself, is
absent, as in the Eunicidae, Syllidae, and Phyllodocidae; or the dorsal bundle may
be absent only in certain regions of the body, as in the hind-body of Terebellids. In
some Amphinomidae and Aphroditidae the notopodium is scarcely distinct as a
separate lobe, being a slight tubercle on the upper surface of the neuropodium;
but the notopodial chaetae are present, and indeed particularly well developed in
many cases.
But whilst, in the Nereidiformia, the parapodia, whether consisting of two lobes or
only one, are always well developed, and project to a more or less pronounced
degree from the sides of the body, it is otherwise in the rest of the group, where
the chaetigerous lobes are usually reduced to mere tubercles or ridges, no doubt
in relation to their burrowing or tubicolous habits. In Sternaspis the chaetae issue
directly from the body-wall.
Amongst the Nereidiformia we find examples in which the parapodia, instead of
being more or less conical "legs," are flattened fore and aft so as to serve as
efficient "fins," as in the active swimmers, Nereis virens and Nephthys caeca, and
in the pelagic Phyllodocids, Alciopids, Typhloscolecids, and Tomopteris.
Of the typical dorsal and ventral cirri, the ventral is only absent in some
Amphinomids amongst the Nereidiformia; the dorsal is absent in Nephthys and
degenerate in Glycera, whilst in a very large number of families of the other sub-
Orders neither cirrus is present. These cirri, though originally filamentous and
sensory, may, by virtue of special blood supply, become "gills," and this occurs in
several families of different sub-Orders. Thus in Eunice this gill is comb-like; in
Amphinome and in Arenicola (on certain segments) it is arborescent, as it is also
in one to three segments in Terebellids; whilst in Ariciidae, Spioniformia,
Cirratulidae, Opheliidae, and Sabellaria it remains more or less finger-shaped or
filamentous. In the family Serpulidae the thoracic cirri, both dorsal and ventral,
become flattened and extended antero-posteriorly, and unite with one another to
form the "thoracic membrane."[320] In Phyllodocidae the cirri are foliaceous and
natatory, and they contain a great quantity of glands of a peculiar character. The
Aphroditidae are distinguished from other Annelids by the possession of "elytra" or
dorsal scales, which appear to be the dorso-ventrally flattened cirri, retaining their
sensory nature, but adding to this function several others.[321]
The chaetae or bristles are mainly used in locomotion, but it is not unreasonable
to believe that some of the stronger, serrated kinds may be used as weapons of
offence and defence; certainly the Polynoids, bristling as they do with stiff chaetae
along each side, must be rather unpleasant to their smaller enemies.
The various bristles may be placed in three chief groups, viz. (1) simple; (2)
jointed; (3) uncini (see Fig. 138).
(1) The simple chaetae may be smooth and hair-shaped, i.e. "capillary," such as
are present in nearly all families: or they may be forked (Amphinomidae), comb-
shaped (Eunice), notched or serrated, or provided with a series of frills at right
angles to their length, as in Aphroditidae; or fringed along one or both sides with a
membranous expansion, as in Terebellids and Sabellids. The simple chaetae may
also be short and spine-like, as in the ventral bundles of Arenicola; or they may be
slightly curved at the end and notched, forming what are generally termed
"crotchets," such as are common amongst Oligochaeta. These "crotchets" may be
simple, or have numerous denticulations at the end (Maldanidae), or be provided
with a membranous hood (Spioniformia, Capitelliformia). In Hermione peculiar
sheathed, spear-like bristles occur (Fig. 138, N).
(2) Jointed chaetae have already been described (p. 246); they are confined to the
sub-Order Nereidiformia, and occur only in certain families.
(3) The uncini are very short chaetae, which are simply embedded in the skin, and
do not extend beyond the body-wall into the body-cavity. An uncinus is a sharply
curved hook, which may have more or less numerous secondary teeth on it. They
are characteristic of the Sabelliformia and the Terebelliformia.
While the chaetae in the Nereidiformia and others are grouped in bundles, those
of many other families are in vertical, transverse rows, as in Maldanidae and in
Arenicola. The uncini are always embedded in such rows, usually slightly raised
from the general level of the body surface, each being termed a "torus
uncinigerus." These tori are usually limited to the sides of the body, but in Myxicola
and in Notomastus they encroach upon the dorsal surface, and in Chaetozone,
also upon the ventral, so as nearly to encircle the body, recalling the
"perichaetous" condition of some earth-worms.
Gills.—We have already seen that several different organs, e.g. the palps in
Sabelliformia, the prostomial tentacles of Chlorhaemidae, and the notopodial cirri
of sundry other Polychaetes, may take on a respiratory function. There are,
however, certain "gills" developed either on the parapodium itself or elsewhere on
the body which it is difficult to homologise. Such are the retractile gills on the
parapodia of the Glyceridae (Fig. 136, C); those of Dasybranchus, near the
abdominal neuropodia; those of Mastobranchus, near the notopodia. Nephthys
has a sickle-shaped gill on the under surface of the notopodium. The long gill
filaments at the posterior end of Sternaspis, again, are only doubtfully interpreted
as the dorsal cirri of some of the posterior segments.
Since primitively the whole skin of the worm is respiratory, any part of the skin may
become more or less specialised for this function, and chiefly, of course, on the
more actively moving parapodia. The blood-vessels constituting the essential part
of the "gill" may make use of any already existing outgrowth (such as a cirrus or a
tentacle), or may push the body-wall out on their own account.
Internal Anatomy.
Probably those organs which have the greatest effect in modifying the shape of
the body are the septa, for we find in the long, free-swimming worms that these
are regularly present throughout the body, and external "segmentation" of the
body is well marked. In burrowing and tubicolous forms the septa are frequently
incompletely developed, or more or fewer may be absent; and the body becomes
less distinctly segmented externally, tends to vary greatly in diameter during
movement, or becomes plumper. With the disappearance of the septa there is also
a diminution in the number of nephridia, as in Arenicola, with only six pairs.
Further, there is frequently a dimorphism of these organs; instead of all of them
serving equally as excretory organs and as genital ducts, some of the most
anterior in the Sabelliformia and Terebelliformia become greatly enlarged, and
take on practically the whole of the former function; whilst more or fewer of the
posterior nephridia dwindle in size, and become genital ducts. The absence of
septa allows a free communication between the successive segments, and thus a
freer flow of coelomic fluid for the distension of the anterior end of the worm during
burrowing.
In the Scoleciformia and Capitelliformia the buccal region exists, but there are no
jaws. In the Sabelliformia and Terebelliformia eversion does not take place and
jaws are absent.
Amongst the Nereidiformia the jaws are absent in the Phyllodocidae and
Hesionidae; when present they are usually set in the direct course of the food.
There may be one small tooth used for stabbing, as in some Syllids (Fig. 141, A);
or a circle of such denticles (Autolytus, Fig. 140, D). To these are added powerful
grasping jaws in Nereis (E); or the latter may alone be present, as in Glycera (F).
In Polynoë the four jaws are carried by hard pieces, to which the muscles are
attached (C and G). In Nephthys there is a dorsal and a ventral jaw.
In the Eunicidae, however, the numerous denticles are carried in a special pouch
below the food tract, with which it communicates anteriorly.[323] They are arranged
in an upper and lower series. The lower series (L) consists of a pair of flat plates
(k) on each side partially embedded in and acted upon by muscles, with a harder
enamelled piece—the actual lower "tooth" (j)—at its anterior end. The upper series
(U) consists of several pieces, varying in shape and size in the various genera of
this family; but developmentally they result from modifications of two rows of small,
similar pieces.[324]
The intestine is generally straight and cylindrical, and is usually constricted by the
septa, if these are present. In the Polynoids the intervening sacculations become
so long as to receive the name of "caeca," which, in Aphrodite, become
enormously elongated (Fig. 142); there are eighteen pairs of them (c), each being
a slender tube bent upon itself, giving off short branches and dilated distally,
where it lies in the base of the parapodium.
Fig. 141.—A, Alimentary canal of Syllid: B, transverse section of pharynx of the
same; b, buccal region; d, oesophageal outgrowth; g, salivary glands; i,
intestine; j, tooth; p, pharynx; s, gizzard: C, alimentary canal of Petta (after
Wirén); i, intestine; o, oesophagus; r, rectum; s, stomach.
Otocysts are rare. Arenicola possesses a pair at the base of the prostomium, each
of which in some species retains an opening to the exterior.[327] They probably
serve as "organs of direction" rather than of "hearing." Aricia and Polyophthalmus
likewise have such organs on the prostomium; whilst Fabricia, Myxicola, Terebella,
and a few others possess them in the peristomium, or in some other segment of
the body.
Fig. 144.—Ammotrypane aulogaster Rathke, enlarged. (From Cuningham.) Anterior
end. a, Prostomium; b, everted buccal region; c, notopodial cirrus; X, ciliated
organ everted; I, II, III, first three segments.
The eggs and spermatozoa in the Polychaeta are discharged into the sea either
by rupture of the body-wall or through the nephridia; the male and female
elements unite, and the resulting fertilised eggs undergo development, either
floating separately in the water, or embedded in jelly, or attached to the body or to
the tube of the worm.
The little animal is thus equipped for an independent life: the provisional chaetae
help in keeping it balanced; and in some cases (Spionidae) serve to protect the
little soft creature, for when it is touched it curls up, and its chaetae stick out at the
sides, so that it looks like a hairy caterpillar. But the larva is quite at the mercy of
the sea, for it is carried hither and thither by currents, and in this way the species
is disseminated. The larvae of the Polychaetes, like those of other animals, occur
at certain periods of the year in large quantities at the surface of the sea, and
serve as food for various larger animals.
These larvae are at first very different from the adult animal, and the necessary
changes to be passed through are more or less great according to the species. It
is not our intention to describe these changes in detail.[329] The larva increases in
size, the permanent chaetae make their appearance in regular order, and the body
exhibits segmentation, the new segments always appearing just in front of the anal
segment. The internal organs gradually develop, and the prostomial and
parapodial appendages grow out in their turn. In the Sabelliformia the
multifilamentous "gills" arise by the continued branching of an at first simple
process (the palp) arising from the latero-ventral surface of each side of the
preoral lobe.[330] These gradually encroach dorsally and ventrally till the
prostomium is more or less encircled; meanwhile the peristomium grows forwards
so as to conceal the prostomium, which no longer increases at the same rate as
does the rest of the body.
Although most worms appear to discharge their ova directly into the sea and take
no further care of them, some make provision for their offspring either by laying
the eggs in a jelly, which will serve as food for the young larvae—Aricia, Ophelia,
Protula, Phyllodoce—or by attaching them to their body. In certain Polynoids the
eggs are attached by means of a secretion to the back, under the elytra, where
they undergo development up to a certain stage. In Exogone and some other
Syllids they are attached to the ventral cirri, or in Grubea limbata, all over the
back. In the female Autolytus (Sacconereis) a ventrally-placed brood sac is formed
by the hardening of a secretion; the eggs develop into embryos inside the brood
sac, and then become free, with head appendages and three pairs of parapodia.
Enormous numbers of such embryos may occur; for instance, some 300 were
counted in a brood sac of Autolytus ebiensis. In the case of tubicolous worms, the
eggs are frequently attached to the tube, either inside or outside. In Spirorbis and
Salmacina the operculum serves as a brood pouch.
Only a very few species are known to be viviparous, viz. Syllis vivipara Kr.,
Cirratulus chrysoderma Clap., Marphysa sanguinea Mont., and Nereis diversicolor
Müll.
In most genera there is no external difference between a mature worm filled with
generative products and an immature one, except, it may be, in the colour; for the
yolk of the eggs is frequently tinted yellow, or pink, or bluish, while the
spermatozoa in mass are white; so that the normal colouring of the worm may be
modified when filled with these elements. But in a few instances striking
anatomical peculiarities are exhibited by the mature worm.[331] In many species of
Nereis, for instance, those segments containing the generative products undergo
more or less extensive changes, while the anterior ones remain unaltered. The
body of the ripe Nereis is then distinguishable into an anterior non-sexual region
and a posterior sexual region; and so great are these changes in certain species
that the mature worms were for a long time believed to belong to a different genus,
and received the name Heteronereis. But we now know their true relations, thanks
to the work of Claparède and others. The males in the Heteronereid phase have
fewer unaltered anterior segments than the females, so that there is a sexual
dimorphism.
The changes which Nereis undergoes in its transformation affect chiefly (a) the
shape of the parapodia, and (b) the form of the chaetae of these parapodia. Other
organs may also be affected, though less noticeably; thus the eyes become
enlarged, the intestine may become so compressed by the generative products as
to be functionless, and the tail develops special sensory papillae.[332]
In the parapodia an increase in size and a sharper delineation of the various parts
take place; then flattened foliaceous outgrowths (Fig. 147, x, y) arise from certain
lobes of the feet, in which, too, the blood supply becomes greatly increased. The
old chaetae are pushed out by the development of new ones of quite a different
shape; these are jointed like the old ones, but the appendix is, in many species at
least, flattened and oar-shaped (Fig. 123, C, p. 246); and the chaetae are
arranged in a fan-like manner. Both these modifications are in evident relation to
the free-swimming habit which the Heteronereid now adopts. The new foot serves
as a swimming organ, the old one was a walking appendage.
But in other genera the hinder genital region of the body becomes separated, on
maturity, from the anterior non-sexual region. Various stages of this "schizogamy,"
or fission into a sexual and a non-sexual zooid, have been observed in different
genera. In the genus Syllis the first segment of the sexual zooid, after its
separation from the asexual zooid, proceeds to bud forth a head. The character of
the head is alike in both sexes, though different species present heads of different
shapes; and as the worms were originally described as distinct genera, the names
then given are retained as descriptive terms. Thus the "Chaetosyllis" form has only
two tentacles; the "Ioda" form has three tentacles and a pair of palps. One and the
same species (e.g. S. hyalina) may successively pass through these stages.
With regard to the asexual portion, there is a regeneration of the tail segments
after the sexual zooid has separated; and the number of segments so regenerated
is usually equal to those that have become sexual. After a time these newly
formed segments will produce generative organs, and take on the characteristic
natatory chaetae, and this region will in its turn separate.
One original "stock," or asexual zooid, thus produces several sexual zooids, but
these are only of one sex for a given stock. The males differ in several important
characters from the females; so different, indeed, are the two sexes that before
their history was worked out by Agassiz[335] they were placed in different genera.
The male zooid has thus come to be known as Polybostrichus (Fig. 149, B). It has
three tentacles and two bifid palps; there are two pairs of peristomial cirri; the
testes are confined to the four anterior segments, which are without natatory
chaetae. The female is termed Sacconereis, owing to the possession of a great
ventral brood sac; its head possesses no separate palps; the peristomium carries
only one cirrus on each side; ova occur in every segment of the body, and may
even extend into the hinder segments of the asexual zooid (Fig. 149, C).
Fig. 149.—Myrianida fasciata. (From Malaquin.) The bright red markings of the living
animal are here represented black. A, An asexual individual which has
produced by budding from the zone (z) a chain of twenty-nine zooids, the
oldest being labelled 1, the youngest 29. B, A ripe male zooid (Polybostrichus),
with three tentacles and a pair of forked palps (p). There are five unaltered
anterior segments. C, A ripe female zooid (Sacconereis) with the palps fused
with the prostomium; s, the ventral brood pouch projecting on each side; t,
tentacles.