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corresponding to those of the maxilla can be clearly traced in the
labium.

Fig. 52.—Maxilla and lower lip of Coleoptera. A, Maxilla of Passalus: a,


cardo; b, stipes; c, palpiger; d, palpus; e, inner or inferior lobe or
lacinia; f, outer or superior lobe or galea: B, Labium of Harpalus
caliginosus: a, mentum; b, hypoglottis; c, palpiger (support of the
labial palp); d, palp; e, ligula; f, paraglossa.

The mentum is an undivided, frequently very hard, piece, continuous


with either the submentum or the gula, and anterior to this are placed
the other parts, viz. the labial palpi and their supports, the palpigers;
beyond and between these exists a central piece (Fig. 52, B, e),
about whose name some difference of opinion prevails, but which
may be called the ligula (languette of French authors), and on each
side of this is a paraglossa. In the Orthoptera the single median
piece—the ligula of Coleopterists—is represented by two divided
parts. In some Insects (many Coleoptera) there is interposed
between the mentum and the palpigers a piece called the hypoglottis
(Fig. 52, B, b). It is not so well ascertained as it should be, that the
pieces of the lower lip bearing the same names in different Orders
are in all cases really homologous, and comparison suggests that
the hypoglottis of Coleoptera may possibly represent the piece
corresponding to the mentum of Orthopterists, the so-called mentum
of beetles being in that case the submentum of Orthopterists.

There is another part of the mouth to which we may call special


attention, as it has recently attracted more attention than it formerly
did; it is a membranous lobe in the interior of the mouth, very
conspicuous in Orthoptera, and called the tongue, lingua, or
hypopharynx; it reposes, in the interior of the mouth (Fig. 51, o), on
the middle parts of the front of the labium; it is probably not entirely
lost in Coleoptera, but enters into the composition of the complex
middle part of the lip by amalgamation with the paraglossae. It has
recently been proposed to treat this lingua as the morphological
equivalent of the labium or of the maxillae, giving it the name of the
endolabium, but the propriety of this course remains to be proved;[20]
the view is apparently suggested chiefly by the structure of the
mouth of Hemimerus, a very rare and most peculiar Insect that has
not as yet been sufficiently studied.

As the maxillae and labium are largely used by taxonomists in the


systematic arrangement of the mandibulate Insects, we give a figure
of them as seen in Coleoptera, where the parts, though closely
amalgamated, can nevertheless be distinguished. This Fig. 52
should be compared with Fig. 51.

In speaking of the segments of the body we pointed out that they


were not separate parts but constituted an uninterrupted whole, and
it is well to remark here that this is also true of the gnathites.
Although the mouth parts are spoken of as separate pieces, they
really form only projections from the great body wall. Fig. 51, B,
shows the intimate connexion that exists between the maxillae and
labium; the continuity of the mandibles with the membrane of the
buccal cavity is capable of very easy demonstration.

The head bears, besides the pieces we have considered, a pair of


antennae. These organs, though varying excessively in form, are
always present in the adult Insect, and exist even in the majority of
young Insects. They are very mobile, highly sensitive organs, situate
on or near the front part of the head. The antennae arise in the
embryo from the procephalic lobes, the morphological import of
which parts is one of the most difficult points connected with Insect
embryology.
The eyes of Insects are of two sorts, simple and compound. The
simple eyes, or ocelli, vary in number from one to as many as
eighteen or twenty; when thus numerous they are situated in groups
on each side of the head. In their most perfect form, as found in adult
aculeate Hymenoptera, in Orthoptera and Diptera, ocelli are usually
two or three in number, and present the appearance of small,
perfectly transparent lenses inserted in the integument. In their
simplest form they are said to consist of some masses of pigment in
connexion with a nerve.

Fig. 53.—Two ommatidia from the eye of Colymbetes fuscus, × 160.


(After Exner.) a, Cornea; b, crystalline cone; c, rhabdom; d,
fenestrate membrane with nerve structures below it; e, iris-
pigment; f, retina-pigment.

The compound, or facetted, eyes are the most remarkable of all the
structures of the Insect, and in the higher and more active forms,
such as the Dragon-flies and hovering Diptera, attain a complexity
and delicacy of organisation that elicit the highest admiration from
every one who studies them. They are totally different in structure
and very distinct in function from the eyes of Vertebrata, and are
seated on very large special lobes of the brain (see Fig. 65), which
indeed are so large and so complex in structure that Insects may be
described as possessing special ocular brains brought into relation
with the lights, shades, and movements of the external world by a
remarkably complex optical apparatus. This instrumental part of the
eye is called the dioptric part in contradistinction from the percipient
portion, and consists of an outer corneal lens (a, Fig. 53), whose
exposed surface forms one of the facets of the eye; under the lens is
placed the crystalline cone (b), this latter being borne on a rod-like
object (c), called the rhabdom. There are two layers of pigment, the
outer (e), called the iris-pigment, the inner (f), the retinal-pigment;
underneath, or rather we should say more central than, the
rhabdoms is the fenestrate membrane (d), beyond which there is an
extremely complex mass of nerve-fibres; nerves also penetrate the
fenestrate membrane, and their distal extremities are connected with
the delicate sheaths by one of which each rhabdom is surrounded,
the combination of sheath and nerves forming a retinula. Each set of
the parts above the fenestrate membrane constitutes an
ommatidium, and there may be many of these ommatidia in an eye;
indeed, it is said that the eye of a small beetle, Mordella, contains as
many as 25,000 ommatidia. As a rule the larvae of Insects with a
complete metamorphosis bear only simple eyes. In the young of
Dragon-flies, as well as of some other Insects having a less perfect
metamorphosis, the compound eyes exist in the early stages, but
they have then an obscure appearance, and are probably
functionally imperfect.

In the interior of the head there exists a horny framework called the
tentorium, whose chief office apparently is to protect the brain. It is
different in kind according to the species. The head shows a
remarkable and unique relation to the following segments. It is the
rule in Insect structure that the back of a segment overlaps the front
part of the one following it; in other words, each segment receives
within it the front of the one behind it. Though this is one of the most
constant features of Insect anatomy, it is departed from in the case
of the head, which may be either received into, or overlapped by, the
segment following it, but never itself overlaps the latter. There is
perhaps but a single Insect (Hypocephalus, an anomalous beetle) in
which the relation between the head and thorax can be considered
to be at all similar to that which exists between each of the other
segments of the body and that following it; and even in
Hypocephalus it is only the posterior angles of the head that overlap
the thorax. Although the head usually appears to be very closely
connected with the thorax, and is very frequently in repose received
to a considerable extent within the latter, it nevertheless enjoys great
freedom of motion; this is obtained by means of a large membrane,
capable of much corrugation, and in which there are seated some
sclerites, so arranged as to fold together and occupy little space
when the head is retracted, but which help to prop and support it
when extended for feeding or other purposes. These pieces are
called the cervical sclerites or plates. They are very largely
developed in Hymenoptera, in many Coleoptera, and in Blattidæ,
and have not yet received from anatomists a sufficient amount of
attention. Huxley suggested that they may be portions of head
segments.

Fig. 54.—Extended head and front of thorax of a beetle, Euchroma: a,


back of head; b, front of pronotum; c, chitinous retractile band; d,
cervical sclerites.

Thorax.

The thorax, being composed of the three consecutive rings behind


the head, falls naturally into three divisions—pro-, meso-, and
metathorax. These three segments differ greatly in their relative
proportions in different Insects, and in different stages of the same
Insect's life. In their more highly developed conditions each of the
three divisions is of complex structure, and the sclerites of which it is
externally made up are sufficiently constant in their numbers and
relative positions to permit of their identification in a vast number of
cases; hence the sclerites have received names, and their
nomenclature is of practical importance, because some, if not all, of
these parts are made use of in the classification of Insects. Each
division of the thorax has an upper region, called synonymically
dorsum, notum, or tergum; an inferior or ventral region, called
sternum; and on each side a lateral region, the pleuron. These
regions of each of the three thoracic divisions are further
distinguished by joining to their name an indication of the segment
spoken of, in the form of the prefixes pro-, meso-, and meta-; thus
the pronotum, prosternum, and propleura make up the prothorax.
The thoracic regions are each made up of sclerites whose
nomenclature is due to Audouin.[21] He considered that every
thoracic ring is composed of the pieces shown in Fig. 55, viz. (1) the
sternum (B', a), an unpaired ventral piece; (2) the notum (A),
composed of four pieces placed in consecutive longitudinal order
(A'), and named praescutum (a), scutum (b), scutellum (c), and post-
scutellum (d); (3) lateral pieces, of which he distinguished on each
side an episternum (B', c), epimeron (e), and parapteron (d), these
together forming the pleuron. We give Audouin's Figure, but we
cannot enter on a full discussion of his views as to the thorax; they
have become widely known, though the constancy of the parts is not
so great as he supposed it would prove to be. Sometimes it is
impossible to find all the elements he thought should be present in a
thoracic ring, while in other cases too many sclerites exist. As a rule
the notum of the meso- and metathoraces is in greater part
composed of two pieces, the scutum and the scutellum; while in the
pronotum only one dorsal piece can be satisfactorily distinguished,
though a study of the development may show that really two are
frequently, if not usually, present. On the other hand, one, or more, of
the notal sclerites in some cases shows evidence of longitudinal
division along the middle. The sternum or ventral piece, though
varying greatly in form, is the most constant element of a thoracic
segment, but it has sometimes the appearance of consisting of two
parts, an anterior and a posterior. The pleuron nearly always
consists quite evidently of two parts, the episternum, the more
anterior and inferior, and the epimeron.[22] The relations between
these two parts vary much; in some cases the episternum is
conspicuously the more anterior, while in others the epimeron is
placed much above it, and may extend nearly as far forwards as it. It
may be said, as a rule, that when the sternum extends farther
backwards than the notum, the epimeron is above the episternum,
as in many Coleoptera; but if the sternum be anterior to the notum,
then the episternum is superior to the epimeron, as in dragon-flies.
We would here again reiterate the fact that these "pieces" are really
not separate parts, but are more or less indurated portions of a
continuous integument, which is frequently entirely occupied by
them; hence a portion of a sclerite that in one species is hard, may in
an allied form be wholly or partly membranous, and in such case its
delimitation may be very evident on some of its sides, and quite
obscure on another.

Fig. 55.—Mesothorax of Dytiscus, after Audouin. A, notum; A', pieces


of the notum separated: a, praescutum; b, scutum; c, scutellum; d,
post-scutellum: B, the sternum and pleura united; B', their parts
separated: a, sternum; c, episternum; d, parapteron; e, epimeron.

The parapteron of Audouin does not appear to be really a distinct


portion of the pleuron; in the case of Dytiscus it is apparently merely
a thickening of an edge. Audouin supposed this part to be specially
connected with the wing-articulation, and the term has been
subsequently used by other writers in connexion with several little
pieces that exist in the pleural region of winged Insects.

The prothorax is even more subject to variation in its development


than the other divisions of the thorax are. In the Hymenoptera the
prosternum is disconnected from the pronotum and is capable,
together with the first pair of legs, of movement independent of its
corresponding dorsal part, the pronotum, which in this Order is
always more or less completely united with the meso-thorax; in the
Diptera the rule is that the three thoracic segments are closely
consolidated into one mass. In the majority of Insects the prothorax
is comparatively free, that is to say, it is not so closely united with the
other two thoracic segments as they are with one another. The three
thoracic rings are seen in a comparatively uniform state of
development in a great number of larvae; also in the adult stages of
some Aptera, and among winged insects in some Neuroptera such
as the Embiidae, Termitidae, and Perlidae. In Lepidoptera the
pronotum bears a pair of erectile processes called patagia; though
frequently of moderately large size, they escape observation, being
covered with scales and usually closely adpressed to the sides of the
pronotum.

The two great divisions of the body—the mesothorax and the


metathorax—are usually very intimately combined in winged Insects,
and even when the prothorax is free, as in Coleoptera, these
posterior two thoracic rings are very greatly amalgamated. In the
higher forms of the Order just mentioned the mesosternum and
mesopleuron become changed in direction, and form as it were a
diaphragm closing the front of the metasternum. The meso- and
meta-thorax frequently each bear a pair of wings.

We have described briefly and figured (Fig. 55) the sclerites of the
mesothorax, and those of the metathorax correspond fairly well with
them. In addition to the sclerites usually described as constituting
these two thoracic divisions, there are some small pieces at the
bases of the wings. Jurine discriminated and named no less than
seven of these at the base of the anterior wing of a Hymenopteron.
One of them becomes of considerable size and importance in the
Order just mentioned, and seems to be articulated so as to exert
pressure on the base of the costa of the wing. This structure attains
its maximum of development in a genus (? nondescript) of Scoliidae,
as shown in Fig. 56. The best name for this sclerite seems to be that
proposed by Kirby and Spence, tegula. Some writers call it
paraptère, hypoptère, or squamule, and others have termed it
patagium; this latter name is, however, inadmissible, as it is applied
to a process of the prothorax we have already alluded to.
Fig. 56.—Head and thorax of wasp from Bogota: t, tegula; b, base of
wing.

To complete our account of the structure of the thorax it is necessary


to mention certain hard parts projecting into its interior, but of which
there is usually little or no trace externally. A large process in many
Insects projects upwards from the sternum in a forked manner. It
was called by Audouin the entothorax; some modern authors prefer
the term apophysis. Longitudinal partitions of very large size,
descending from the dorsum into the interior, also exist; these are
called phragmas, and are of great importance in some Insects with
perfect flight, such as Hymenoptera, Lepidoptera, and Diptera. There
is no phragma in connection with the pronotum, but behind this part
there may be three. A phragma has the appearance of being a fold
of the dorsum; it serves as an attachment for muscles, and may
probably be of service in other ways. More insignificant projections
into the interior are the little pieces called apodemes (Fig. 57, e);
these are placed at the sides of the thorax near the wings. The
apophyses are no doubt useful in preserving the delicate vital organs
from shocks, or from derangement by the muscular movements and
the changes of position of the body.

Fig. 57.—Transverse section of skeleton of metathorax of Goliathus


druryi, seen from behind: a, metanotum; b, metasternum; c,
phragma; d, entothorax (apophysis or furca); e, apodeme; f,
tendon of articulation. (After Kolbe.)

The appendages of the thorax are (a) inferior, the legs; (b) superior,
the wings. The legs are always six in number, and are usually
present even in larvae, though there exist many apodal larvae,
especially in Diptera. The three pairs of legs form one of the most
constant of the characters of Insects. They are jointed appendages
and consist of foot, otherwise tarsus; tibia, femur, trochanter, and
coxa; another piece, called trochantin more or less distinctly
separated from the coxa, exists in many Insects. The legs are
prolongations of the body sac, and are in closer relation with the
epimera and with the episterna than with other parts of the crust,
though they have a close relation with the sternum. If we look at the
body and leg of a neuropterous Insect (Fig. 58) we see that the basal
part of the leg—the coxa—is apparently a continuation of one of the
two pleural pieces or of both; in the latter case one of the prolonged
pieces forms the coxa proper, and the tip of the other forms a
supporting piece, which may possibly be the homologue of the
trochantin of some Insects. In some Orthoptera, especially in
Blattidae, and in Termitidae, there is a transverse chitinised fold
interposed between the sternum and the coxa, and this has the
appearance of being the same piece as the trochantin of the anterior
legs of Coleoptera.

Fig. 58.—Hind leg of Panorpa: a, episternum; a′, epimeron; b, coxa; b′,


coxal fold of epimeron; c, trochanter; d, femur; e, tibia; f, tarsus.

Beyond the coxa comes the trochanter; this in many Hymenoptera is


a double piece, though in other Insects it is single; usually it is the
most insignificant part of the leg. The femur is, on the whole, the
least variable part of the leg; the tibia, which follows it, being
frequently highly modified for industrial or other purposes. The joint
between the femur and the tibia is usually bent, and is therefore the
most conspicuous one in the leg; it is called the knee. The other
joints have not corresponding names, though that between the tibia
and the tarsus is of great importance. The spines at the tip of the
tibia, projecting beyond it, are called spurs, or calcares. The tarsus
or foot is extremely variable; it is very rarely absent, but may consist
of only one piece—joint, as it is frequently called[23]—or of any larger
number up to five, which may be considered the characteristic
number in the higher Insect forms. The terminal joint of the tarsus
bears normally a pair of claws; between the claws there is frequently
a lobe or process, according to circumstances very varied in different
Insects, called empodium, arolium, palmula, plantula,
pseudonychium, or pulvillus. This latter name should only be used in
those cases in which the sole of the foot is covered with a dense
pubescence. The form of the individual tarsal joints and the armature
or vestiture of the lower surface are highly variable. The most
remarkable tarsus is that found on the front foot of the male
Dytiscus.

It has been suggested that the claws and the terminal appendage of
the tarsus ought to be counted as forming a distinct joint; hence
some authors state that the higher Insects have six joints to the feet.
These parts, however, are never counted as separate joints by
systematic entomologists, and it has recently been stated that they
are not such originally.

The parts of the foot at the extremity of the last tarsal joint proper are
of great importance to the creature, and vary greatly in different
Insects. The most constant part of this apparatus is a pair of claws,
or a single claw. Between the two claws there may exist the
additional apparatus referred to above. This in some Insects—
notably in the Diptera—reaches a very complex development. We
figure these structures in Pelopaeus spinolae, a fossorial
Hymenopteron, remarking that our figures exhibit the apparatus in a
state of retraction (Fig. 59). According to the nomenclature of Dahl
and Ockler[24] the plate (b) on the dorsal aspect is the pressure plate
(Druck-Platte), and acts as an agent of pressure on the sole of the
pad (C, e); c and d on the underside are considered to be extension-
agents; c, extension-plate; d, extension-sole (Streck-Platte, Streck-
Sohle). These agents are assisted in acting on the pad by means of
an elastic bow placed in the interior of the latter. The pad (e) is a
very remarkable structure, capable of much extension and retraction;
when extended it is seen that the pressure plate is bent twice at a
right angle so as to form a step, the distal part of which runs along
the upper face of the basal part of the pad; the apical portion of this
latter consists of two large lobes, which in repose, as shown in our
Figure (f), fall back on the pad, something in the fashion of the
retracted claws of the cat, and conceal the pressure-plate.

The mode in which Insects are able to walk on smooth perpendicular


surfaces has been much discussed, and it appears highly probable
that the method by which this is accomplished is the exudation of
moisture from the foot; there is still, however, much to be ascertained
before the process can be satisfactorily comprehended. The theory
to the effect that the method is the pressure of the atmosphere
acting on the foot when the sole is in perfect apposition with the
object walked on, or when a slight vacuum is created between the
two, has apparently less to support it.

Fig. 59.—Foot of Pelopaeus, a fossorial wasp: A, tarsus entire; B,


terminal joint, upper side; C, under side. a, claw; b, base of
pressure-plate; c, extension-plate; d, extension-sole; e, pad; f,
lobe of pad retracted.

The legs of the young Insect are usually more simple than those of
the adult, and in caterpillars they are short appendages, and only
imperfectly jointed. If a young larva, with feet, of a beetle, such as
Crioceris asparagi be examined, it may be seen that the leg is
formed by protuberance of the integument, which becomes divided
into parts by simple creases; an observation suggesting that the
more highly developed jointed leg is formed in a similar manner. This
appears to be really the case, for the actual continuity of the limb at
the chief joint—the knee—can be demonstrated in many Insects by
splitting the outer integument longitudinally and then pulling the
pieces a little apart; while in other cases even this is not necessary,
the knee along its inner face being membranous to a considerable
extent, and the membrane continuous from femur to tibia.

Turning to the wings, we remark that there may be one or two pairs
of these appendages. When there is but one pair it is nearly always
mesothoracic, when there are two pairs one is invariably
mesothoracic, the other metathoracic. The situation of the wing is
always at the edge of the notum, but the attachment varies in other
respects. It may be limited to a small spot, and this is usually the
case with the anterior wing; or the attachment may extend for a
considerable distance along the edge of the notum, a condition
which frequently occurs, especially in the case of the posterior
wings. The actual connexion of the wings with the thorax takes place
by means of strong horny lines in them which come into very close
relation with the little pieces in the thorax which we have already
described, and which were styled by Audouin articulatory epidemes.
There is extreme variety in the size, form, texture, and clothing of the
wings, but there is so much resemblance in general characters
amongst the members of each one of the Orders, that it is usually
possible for an expert, seeing only a wing, to say with certainty what
Order of Insects its possessor belonged to. We shall allude to these
characters in treating of the Orders of Insects.

Each wing consists of two layers, an upper and a lower, and


between them there may be tracheae and other structures,
especially obvious when the wings are newly developed. It has been
shown by Hagen that the two layers can be separated when the
wings are recently formed, and it is then seen that each layer is
traversed by lines of harder matter, the nervures. These ribs are
frequently called wing-veins, or nerves, but as they have no relation
to the anatomical structures bearing those names, it is better to
make use of the term nervures. The strength, number, form and
inter-relations of these nervures vary exceedingly; they are thus
most important aids in the classification of Insects. Hence various
efforts have been made to establish a system of nomenclature that
shall be uniform throughout the different Orders, but at present
success has not attended these efforts, and it is probable that no
real homology exists between the nervures of the different Orders of
Insects. We shall not therefore discuss the question here. We may,
however, mention that German savants have recently distinguished
two forms of nervures which they consider essentially distinct, viz.
convex and concave. These, to some extent, alternate with one
another, but a fork given off by a convex one is not considered to be
a concave one. The terms convex and concave are not happily
chosen; they do not refer to the shape of the nervures, but appear to
have been suggested by the fact that the surface of the wing being
somewhat undulating the convex veins more usually run along the
ridges, the concave veins along the depressions. The convex are the
more important of the two, being the stronger, and more closely
connected with the articulation of the wing.

The wings, broadly speaking, may be said to be three-margined: the


margin that is anterior when the wings are extended is called the
costa, and the edge that is then most distant from the body is the
outer margin, while the limit that lies along the body when the wings
are closed is the inner margin.

The only great Order of Insects provided with a single pair of wings
is the Diptera, and in these the metathorax possesses, instead of
wings, a pair of little capitate bodies called halteres or poisers. In the
abnormal Strepsiptera, where a large pair of wings is placed on the
metathorax, there are on the mesothorax some small appendages
that are considered to represent the anterior wings. In the great
Order Coleoptera, or beetles, the anterior wings are replaced by a
pair of horny sheaths that close together over the back of the Insect,
concealing the hind-wings, so that the beetle looks like a wingless
Insect: in other four-winged Insects it is usually the front wings that
are most useful in flight, but the elytra, as these parts are called in
Coleoptera, take no active part in flight, and it has been recently
suggested by Hoffbauer[25] that they are not the homologues of the
front wings, but of the tegulae (see Fig. 56), of other Insects. In the
Orthoptera the front wings also differ in consistence from the other
pair over which they lie in repose, and are called tegmina. There are
many Insects in which the wings exist in a more or less rudimentary
or vestigial condition, though they are never used for purposes of
flight.

The abdomen, or hind body, is the least modified part of the body,
though some of the numerous rings of which it is composed may be
extremely altered from the usual simple form. Such change takes
place at its two extremities, but usually to a much greater extent at
the distal extremity than at the base. This latter part is attached to
the thorax, and it is a curious fact that in many Insects the base of
the abdomen is so closely connected with the thorax that it has all
the appearance of being a portion of this latter division of the body;
indeed it is sometimes difficult to trace the real division between the
two parts. In such cases a further differentiation may occur, and the
part of the abdomen that on its anterior aspect is intimately attached
to the thorax may on its posterior aspect be very slightly connected
with the rest of the abdomen. Under such circumstances it is difficult
at first sight to recognise the real state of the case. When a segment
is thus transferred from the abdomen to the metathorax, the part is
called a median segment. The most remarkable median segment
exists in those Hymenoptera which have a stalked abdomen, but a
similar though less perfect condition exists in many Insects. When
such a union occurs, it is usually most complete on the dorsal
surface, and the first ventral plate may almost totally disappear: such
an alteration may involve a certain amount of change in the sclerites
of the next segment, so that the morphological determination of the
parts at the back of the thorax and front of the abdomen is by no
means a simple matter. A highly modified hind-body exists in the
higher ants, Myrmicidae. In Fig. 60 we contrast the simple abdomen
of Japyx with the highly modified state of the same part in an ant.
Fig. 60.—Simple abdomen of Japyx (A) contrasted with the highly
modified one of an ant, Cryptocerus (B). The segments are
numbered from before backwards.

Unlike the head and thorax, the abdomen is so loosely knitted


together that it can undergo much expansion and contraction. This is
facilitated by an imbricated arrangement of the plates, and by their
being connected by means of membranes admitting of much
movement (Fig. 47, m, p. 88). In order to understand the structure of
the abdomen it should be studied in its most distended state; it is
then seen that there is a dorsal and a ventral hard plate to each ring,
and there is also usually a stigma; there may be foldings or plications
near the line of junction of the dorsal and ventral plates, but these
margins are not really distinct pieces. The pleura, in fact, remain
membranous in the abdominal region, contrasting strongly with the
condition of these parts in the thorax. The proportions of the plates
vary greatly; sometimes the ventral are very large in proportion to the
dorsal, as is usually the case in Coleoptera, while in the Orthoptera
the reverse condition prevails.

Cerci or other appendages frequently exist at the extremity of the


abdomen (Fig. 47, n, p. 88); the former are sometimes like antennae,
while in other cases they may be short compressed processes
consisting of very few joints. The females of many Insects possess
saws or piercing instruments concealed within the apical part of the
abdomen; in other cases an elongate exserted organ, called
ovipositor, used for placing the eggs in suitable positions, is present.
Such organs consist, it is thought, either of modified appendages,
called gonapophyses, or of dorsal, ventral, or pleural plates. The
males frequently bear within the extremity of the body a more or less
complicated apparatus called the genital armour. The term
gonapophysis is at present a vague one, including stings, some
ovipositors, portions of male copulatory apparatus, or other
structures, of which the origin is more or less obscure.

The caterpillar, or larva, of the Lepidoptera and some other Insects,


bears a greater number of legs than the three pairs we have
mentioned as being the normal number in Insects, but the posterior
feet are in this case very different from the anterior, and are called
false legs or prolegs. These prolegs, which are placed on the hind
body, bear a series of hooks in Lepidopterous larvae, but the
analogous structures of Sawfly larvae are destitute of such hooks.

Placed along the sides of the body, usually quite visible in the larva,
but more or less concealed in the perfect Insect, are little apertures
for the admittance of air to the respiratory system. They are called
spiracles or stigmata. There is extreme variety in their structure and
size; the largest and most remarkable are found on the prothorax of
Coleoptera, especially in the groups Copridae and Cerambycidae.

The exact position of the stigmata varies greatly, as does also their
number. In the Order Aptera there may be none, while the maximum
number of eleven pairs is said by Grassi[26] to be attained in Japyx
solifugus: in no other Insect have more than ten pairs been recorded,
and this number is comparatively rare. Both position and number
frequently differ in the early and later stages of the same Insect. The
structure of the stigmata is quite as inconstant as the other points we
have mentioned are.

Fig. 61.—Membranous space between pro- and meso-thoraces of a


beetle Euchroma, showing stigma (st); a, hind margin of
pronotum; b, front leg; c, front margin of mesonotum; d, base of
elytra; e, mesosternum.
The admission of air to the tracheal system and its confinement
there, as well as the exclusion of foreign bodies, have to be provided
for. The control of the air within the system is, according to
Landois[27] and Krancher,[28] usually accomplished by means of an
occluding apparatus placed on the tracheal trunk a little inside of the
stigma, and in such case this latter orifice serves chiefly as a means
for preventing the intrusion of foreign bodies. The occluding
apparatus consists of muscular and mechanical parts, which differ
much in their details in different Insects. Lowne supposes that the air
is maintained in the tracheal system in a compressed condition, and
if this be so, this apparatus must be of great importance in the Insect
economy. Miall and Denny[29] state that in the anterior stigmata of
the cockroach the valves act as the occluding agents, muscles being
attached directly to the inner face of the valves, and in some other
Insects the spiracular valves appear to act partially by muscular
agency, but there are many stigmata having valves destitute of
muscles. According to Lowne[30] there exist valves in the blowfly at
the entrance to the trachea proper, and he gives the following as the
arrangement of parts for the admission of air:—there is a spiracle
leading into a chamber, the atrium, which is limited inwardly by the
occluding apparatus; and beyond this there is a second chamber, the
vestibule, separated from the tracheae proper by a valvular
arrangement. He considers that the vestibule acts as a pump to
force the air into the tracheae.

Fig. 62.—Diagrammatic Insect to explain terms of position. A, apex; B,


base: 1, tibia; 2, last abdominal segment; 3, ideal centre.

Systematic Orientation.
Terms relating to position are unfortunately used by writers on
entomology in various, even in opposite senses. Great confusion
exists as to the application of such words as base, apex, transverse,
longitudinal. We can best explain the way in which the relative
positions and directions of parts should be described by reference to
Figure 62. The spot 3 represents an imaginary centre, situated
between the thorax and abdomen, to which all the parts of the body
are supposed to be related. The Insect should always be described
as if it were in the position shown in the Figure, and the terms used
should not vary as the position is changed. The creature is placed
with ventral surface beneath, and with the appendages extended,
like the Insect itself, in a horizontal plane. In the Figure the legs are,
for clearness, made to radiate, but in the proper position the anterior
pair should be approximate in front, and the middle and hind pairs
directed backwards under the body. The legs are not to be treated as
if they were hanging from the body, though that is the position they
frequently actually assume. The right and left sides, and the upper
and lower faces (these latter are frequently also spoken of as sides),
are still to retain the same nomenclature even when the position of
the specimen is reversed. The base of an organ is that margin that is
nearest to the ideal centre, the apex that which is most distant. Thus
in Fig. 62, where 1 indicates the front tibia, the apex (A) is broader
than the base (B); in the antennae the apex is the front part, while in
the cerci the apex is the posterior part; in the last abdominal
segment (2) the base (B) is in front of the apex (A). The terms
longitudinal and transverse should always be used with reference to
the two chief axes of the body-surface; longitudinal referring to the
axis extending from before backwards, and transverse to that going
across, i.e. from side to side.

CHAPTER IV

ARRANGEMENT OF INTERNAL ORGANS–MUSCLES–NERVOUS SYSTEM–


GANGLIONIC CHAIN–BRAIN–SENSE-ORGANS–ALIMENTARY CANAL–
MALPIGHIAN TUBES–RESPIRATION–TRACHEAL SYSTEM–FUNCTION OF
RESPIRATION–BLOOD OR BLOOD-CHYLE–DORSAL VESSEL OR HEART–
FAT-BODY–OVARIES–TESTES–PARTHENOGENESIS–GLANDS.

The internal anatomy of Insects may be conveniently dealt with


under the following heads:—(1) Muscular system; (2) nervous
system; (3) alimentary system (under which may be included
secretion and excretion, about which in Insects very little is known);
(4) respiratory organs; (5) circulatory system; (6) fat-body; (7)
reproductive system.

Fig. 63.—Diagram of arrangement of some of the internal organs of an


Insect: a, mouth; b, mandible; c, pharynx; d, oesophagus; e,
salivary glands (usually extending further backwards); f, eye; g,
supra-oesophageal ganglion; h, sub-oesophageal ganglion; i,
tentorium; j, aorta; k1, k2, k3, entothorax; l1-l8, ventral nervous
chain; m, crop; n, proventriculus; o, stomach; p, Malpighian tubes;
q, small intestine; r, large intestine; s, heart; t, pericardial septum;
u, ovary composed of four egg-tubes; v, oviduct; w, spermatheca
(or an accessory gland); x, retractile ovipositor; y, cercus; z,
labrum.

Many of the anatomical structures have positions in the body that are
fairly constant throughout the class. Parts of the respiratory and
muscular systems and the fat-body occur in most of the districts of
the body. The heart is placed just below the dorsal surface; the
alimentary canal extends along the middle from the head to the end
of the body. The chief parts of the nervous system are below the
alimentary canal, except that the brain is placed above the beginning
of the canal in the head. The reproductive system extends in the
abdomen obliquely from above downwards, commencing anteriorly

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