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211) is not a pupa obtecta, but has the head and appendages free,
and is provided with enormous mandibles. Although these Insects in
general appearance resemble Eriocephala to such an extent that
both have been placed in one genus, viz. Micropteryx, yet the two
forms are radically distinct. The most remarkable point in Micropteryx
is the metamorphosis; the female moth is furnished with a cutting
ovipositor, by the aid of which she deposits an egg between the two
layers of a leaf after the manner of a saw-fly;[342] the larva mines the
newly-opened leaves in the early spring, and feeds up with rapidity; it
by some means reaches the ground, and there pupates in a firm but
thin cocoon, with grains of earth fastened to it; in this it passes the
greater part of its life as a larva, changing to a pupa very early in the
following spring. The pupa is unlike any other Lepidopterous pupa,
but is similar to those of Trichoptera; neither the head nor the
appendages are glued to the body or to one another, but are free, so
that the pupa can use the appendages to a considerable extent; it is
furnished with enormous mandibles (Fig. 211, C, D), which are
detached and shed after emergence.[343] In the interval between the
larval period of feeding and the imaginal instar, the phenomena of
life are essentially like those of Trichoptera. The larva has not been
at all satisfactorily studied; the spiracles appear to be excessively
minute, but have been ascertained by Dr. Chapman to be normal in
number and position.

All the information we possess points to profound distinctions

between Micropteryx and Eriocephala, for whereas in the former the
mandibles drop off from the pupa, so that the imago has no
mandibles, in the latter the mandibles exist, as they do in several
other true Lepidoptera. As the history of the mandibles is not known
in other Lepidoptera (where they are present in the larva but wanting
in the imago), it is premature to conclude that no other Lepidoptera
suffer the actual loss of the mandibles as Micropteryx does, though
there is nothing to lead us to believe that in any other Lepidopterous
pupa are the mandibles specially developed as they are in
Micropteryx. This pupa is in fact quite unique in this Order of Insects.
When the history of the pupal mandibles is known, we shall be able
to decide whether they are secondary structures, like the deciduous,
supplementary mandibles found in Otiorhynchides (Coleoptera,
Rhynchophora).
 CHAPTER VII

DIPTERA—OR FLIES; APHANIPTERA—OR FLEAS; THYSANOPTERA—OR

THRIPS

Order VII. Diptera

Wings two, membranous, usually transparent and never very

large; behind the wings a pair of small erect capitate bodies—
halteres—frequently concealed under membranous hoods. No
distinct prothorax, all the divisions of the thorax being united to
form a large mass. Mouth-parts very variable, formed for suction
not for biting, frequently assuming the form of a proboscis that
can be retracted and concealed in a cleft of the under side of the
head. The metamorphosis is very great, the larvae bearing no
resemblance whatever to the perfect Insects, but being usually
footless grubs or maggots; frequently the head is indistinct,
small, and retracted. Pupa variable, either exposed and rather
hard, with the appendages of the body more or less adherent; or
enclosed in a scaly capsule looking like a seed, and when
extracted, soft and delicate, with the appendages not fastened to
the body incapable of movement.

This definition of the Diptera, or two-winged flies, is framed without

reference to the fleas, which are wingless, or to a few other parasitic
wingless Diptera, such as the sheep-tick. Although the Order is of
enormous extent, these exceptional cases are remarkably few.
About 40,000 species of Diptera have been discovered, but these
are only a tithe of what are still unknown to science. The Order is not
a favourite one with entomologists, and by the rest of the world it
may be said to be detested. Flies do not display the sort of
intelligence we appreciate, or the kind of beauty we admire, and as a
few of the creatures somewhat annoy us, the whole Order is only too
frequently included in the category of nuisances that we must submit
to. Moreover, the scavenger-habits that are revealed, when we begin
to study their lives, are very repugnant to many persons. It is
therefore no wonder that flies are not popular, and that few are
willing to study them, or to collect them for observation.
Nevertheless, Diptera have considerable claims to be classed as
actually the highest of Insects physiologically, for it is certainly in
them that the processes of a complete life-history are carried on with
the greatest rapidity and that the phenomena of metamorphosis
have been most perfected. A maggot, hatching from an egg, is able
to grow with such rapidity that the work of its life in this respect is
completed in a few days; then forming an impenetrable skin it
dissolves itself almost completely; solidifying subsequently to a sort
of jelly, it in a few days reconstructs itself as a being of totally
different appearance and habits, in all its structures so profoundly
changed from what it was that the resources of science are severely
taxed to demonstrate any identity of the organs of the two instars.
 Fig. 212—A Dipteron (Fam. Syrphidae), Cheilosia chrysocoma. Britain.
A, Adult larva; B, the pupa; C, nymph, extracted from pupa; D,
imago. (From Weyenbergh.)

A good study of the comparative anatomy of Diptera has never been

made; Baron Osten Sacken, one of our most accomplished
Dipterologists, has recently stated that "the external characters of
the Diptera have as yet been very insufficiently studied." We shall
therefore only trouble the student with a few observations on points
of structure that are of special importance, or that he will find
frequently alluded to. The head is remarkable for its mobility, and is
connected with the thorax by a slender concealed neck that permits
the head to undergo semi-rotation. A large part—sometimes nearly
the whole—of the exposed surface of the head is occupied by the
faceted eyes. It is usually the case that the eyes are larger in the
male than in the female, and the sexual discrepancy in this respect
may be very great. When the eyes of the two sides meet in a
coadapted line of union the Insect is said to be "holoptic," and when
the eyes are well separated "dichoptic."[344] The holoptic condition is
specially characteristic of the male, but in some forms occurs in both
sexes. There is no definite distinction between holoptic and dichoptic
eyes. The eyes may be enormous, Fig. 238, without actually uniting,
and in the cases where actual contiguity occurs, it takes place in
different manners.[345] The eyes are frequently during life of brilliant
colours and variegate with stripes or spots; this condition disappears
speedily after death, and it is uncertain what the use of this
coloration may be.[346] The eyes are frequently densely set with
hairs between the almost innumerable facets. These facets
frequently differ in size according to their position in the organ. The
curious double eye of the male Bibio (cf. Fig. 224) is well worth
notice. There are usually three small ocelli placed very near together
on the middle of the summit of the head.

The antennae are of considerable importance, as they offer one of

the readiest means of classification. The families placed by
systematists at the commencement of the Order have antennae
similar to those of the majority of Insects, inasmuch as they consist
of a series of segments approximately similar to one another, and
arranged in a linear manner (Fig. 213, A). The number of these joints
is never very great, but reaches sixteen in certain Tipulidae, and falls
as low as eight in some Bibionidae. In certain cases where the
antennae of the male are densely feathered (Chironomus, e.g.), the
number of joints is in that sex greatly augmented, but they are
imperfectly separated. This form of antenna gives the name
Nemocera to the first series of Diptera.

Fig. 213—Antennae of flies. A, The two antennae of Glaphyroptera

picta (Mycetophilidae); B, antenna of Hexatoma pellucens
(Tabanidae); C, of Asilus crabroniformis (Asilidae); D, of Leptis
scolopacea (Leptidae); E, of Dolichopus undulatus (Dolichopidae);
F, of Volucella bombylans (Syrphidae). (After Wandolleck.)

The majority of flies have antennae of another form, peculiar to the

Order, viz. three segments, the outer one of which is of diverse form,
according to the genus or species, and bears on its front a fine
projecting bristle, frequently feathered, as in Fig. 213, F; and often
distinctly divided into two or more joints. This form of antenna is
found in the series Aschiza and Schizophora; it is well exemplified in
the common house-fly, where the organs in question hang from the
forehead, and are placed in a hollow formed for their reception on
the front of the head. Flies with this form of antennae are called
Athericerous. Between the two forms of antennae we have
mentioned there exists what may, speaking roughly, be called an
intermediate condition, or rather a variety of intermediate conditions,
associated in the series Brachycera (Fig. 213, B to D).[347] Here
there are three (sometimes one or two) segments and a terminal
appendage, but the appendage is usually compound (often so
distinctly compound that it is evidently a series of partially, or even
completely, separate joints, Fig. 213, B): the appendage in these
cases is terminal, that is to say it is placed, not as in the Eumyiidae
on the front of the joint that bears it, but (in the great majority of
Brachycera) at the tip thereof; this appendage is often conical and
pointed, often hair-like. Exceptional forms of antenna are found in
the parasitic flies of the series Pupipara. In the Order generally the
two basal joints of the antennae are evidently distinct in function from
the others, and form the "scape"; the part of the antenna beyond the
scape is called the "flagellum"; an appendage of the flagellum is
called "arista" when bristle-like, when thicker "style." In the basal
joint of the antenna there is a complex nervous structure known as
Johnston's organ. It is specially well developed in Culex and
Chironomus, and is larger in the male than it is in the female. Child
has found something of the kind present in all the Diptera he has
examined, and he considers that an analogous structure exists in
Insects of other Orders. He thinks it is concerned with the perception
of vibration, there being no sharp distinction between auditory and
tactile sensation.[348]

About one-half of the Diptera possess a peculiar structure in the form

of a head-vesicle called "ptilinum." In the fly emerging from the pupa
this appears as a bladder-like expansion of the front of the head;
being susceptible of great distension, it is useful in rupturing the hard
shell in which the creature is then enclosed. In the mature fly the
ptilinum is completely introverted, and can be found only by
dissection; a little space, the "lunula," just under an arched suture,
extending over the point of insertion of the antennae remains,
however, and offers evidence of the existence of the ptilinum. This
structure is also of importance in classification, though, unfortunately,
it is difficult to verify.[349]
No point of Insect morphology has given rise to more difference of
opinion than the mouth of Diptera; and the subject is still very far
from being completely understood. The anatomy and morphology of
the mandibulate Insect-mouth are comparatively simple (though not
without greater difficulties than are usually appreciated); and it has
been the desire of morphologists to homologise the sucking mouth of
Diptera with the biting mouth; hence the view that the appendages of
three segments are separate and distinct in the fly's mouth is taken
for granted, and it is further assumed that some of the secondary
parts of the appendages of the biting mouth can also be recognised
in the sucking mouth. The anatomy of the mouth-parts is, however,
subject to great diversity of structure within the limits of the Order
itself, even the two sexes in some species differing profoundly in this
respect.[350] In the majority of the family Oestridae the mouth-parts
are practically absent, and no definite entry to the alimentary canal
can be perceived (Fig. 245). Besides this condition and its antithesis
(Fig. 214), the complex assemblage of lancets seen in the Breeze-
flies that draw blood, there is a great variety of other anatomical
conditions.

Fig. 214.—Mouth-parts of the common blood-sucking fly, Haematopota

pluvialis ♀ . A, Viewed from beneath, the proboscis removed; a,
labrum; b, b, cultelli (mandibles of other anatomists); c, c,
scalpella (maxillae of other anatomists); d, part of ventral scutum
of second metamere; e, e, f, f, parts of palpi; g, hypopharynx and
pellucid salivary duct; h, salivary receptacle; i, salivary duct; k,
membranous part of second metamere; l, pharynx: B, labrum,
pharynx, hypopharynx, separated, seen from beneath; a, labrum;
b, hypopharynx; c, salivary duct; d, pharynx; e, protractor
muscles: C, proboscis (labium) from beneath; a, scutum
proboscidis; c, c, labella; d, d, retractor muscles. (After Meinert).
Although, as we have said, great diversity of opinion exists, yet on
the whole the majority of Dipterologists accept a view something to
the following effect:—the labrum, or the labrum combined with the
epipharynx, is frequently much prolonged; the tongue—hypopharynx
—may also be much prolonged, and may form, in apposition with the
labrum, a more or less imperfect tube for ingestion of the nutriment;
the labium is more or less membranous or fleshy, and acts as a
sheathing organ, its tips—called labella—-being in some cases
developed to a quite extraordinary extent. As to the other parts of the
mouth there is less agreement; the pointed organs (Fig. 214, A, b b)
are by many identified as mandibles, while another pair of pointed
processes (c c) are considered to be parts of a maxilla, and the palpi
(f f) are by some considered to be maxillary palps. The Danish
entomologist, Meinert, has published the best anatomical description
of many of the diverse kinds of Dipterous mouth.[351] He, however,
takes a different view of the morphology; he considers that not only
may parts of the appendages of the mouth be much modified during
the early stages of the individual development, but that they may be
differently combined, even parts of the appendages of two segments
being brought together in intimate combination. He has also pointed
out that the mandibulate and sucking mouth are mechanical
implements constructed on opposed principles; the main object of a
biting mouth being the fixing and perfecting of the articulations of the
mouth, so that great power of holding may be attained with a limited
but definite power of movement. In the sucking mouth the parts are
intimately associated for simple protrusion. Hence the two kinds of
mouth must have been distinguished very early in the phylogeny, so
that we must not expect to find a great correspondence between the
parts of biting and sucking mouths. He apparently also considers
that not only the appendages of a head-segment, but also part of the
body of the segment, may be used in the construction of the mouth-
organs. Meinert's views allow a much greater latitude of
interpretation of the parts of the Dipterous mouth; had he contented
himself with enunciating them in the manner we have followed him in
summarily describing, they would have been recognised as a
formidable obstacle to the facile adoption of the ordinary views. He
has, however, accompanied his general statement with a particular
interpretation and a distinct nomenclature, neither of which is it
possible to adopt at present, as they have no more justification than
the ordinary view. So that instead of one set of doubtful
interpretations we have two.[352] In so difficult a question as
homologising the trophi of different Orders of Insects we ought to
use exhaustively every method of inquiry: and from this point of view
the development is of great importance. This has, however, as yet
thrown but little light on the subject, this study being a very difficult
one owing to the profound changes that take place during
metamorphosis, the diversity of the parts in the early stages of
Diptera, and the possibility that the larval conditions may themselves
have been greatly changed in the course of the phylogeny. Miall
informs us, however, that in Chironomus as well as in Corethra the
new parts of the mouth of the imago are developed within those of
the larva.[353] This may permit of an identification of the main
divisions of the mouth, at any rate in these cases. Lowne has to
some extent traced the development in the blowfly, and he does not
agree with the usual interpretation of the parts in the adult.

The mouth is of considerable importance in the classification of

Diptera. The Nemocera are remarkable from the linear development
and flexibility of the palpi, which are nearly always at least three- or
four-jointed; this condition occurring in no other Diptera. The palpi
attain an extraordinary development in some Culicidae; in the genus
Megarrhina they are nearly as long as the body, and project in front
of the head after the fashion of the palpi of Lepidoptera. In the
Brachycera the sclerites or hard parts of the mouth reach a
maximum of development, and in Tabanidae (Fig. 214),
Nemestrinidae and Bombyliidae are often quite disproportionate to
the size of the Insect. In many of the Eumyiid flies the soft parts are
greatly developed, and capable of a variety of movement, the
proboscis as a whole being protrusible, and having an elbow-joint in
the middle.

The thorax is remarkable from the absence of distinct separation into

the three divisions that may usually be so easily distinguished in
Insects. The perfect combination of the three segments adds much
to the difficulty of arriving at general conclusions as to the
identification of the parts; hence considerable difference of opinion
still prevails. It was formerly supposed that a segment from the
abdomen was added to the thorax of Diptera as it is in Hymenoptera,
but this has been shown by Brauer to be erroneous. Indeed,
according to Lowne, the abdominal cavity is increased by the
addition of the small posterior area of the thorax; it being the
mesophragma that separates the second and third great divisions of
the body-cavity. The prothorax is always small, except in a few of the
abnormal wingless forms (Melophagus); in Nycteribia (Fig. 248) the
mesothorax forms the anterior part of the body; the head and such
parts of the prothorax as may be subsequently discovered to exist
being placed entirely on the dorsum of the body. The mesothorax in
all the winged Diptera forms by far the larger portion of the thoracic
mass, the prominent part of it, that projects backwards to a greater
or less extent over the base of the abdomen, being the scutellum.
The first or prothoracic stigma is remarkably large and distinct, and is
by some called mesothoracic. Another large stigma is placed very
near to the halter (or balancer); the metathorax being very small. An
imperfect stigma is said by Lowne to exist in the blowfly near the
base of the wing. The number of abdominal segments externally
visible is very diverse; there may be as many as nine (in the male
Tipula), or as few as five, or even four, when the basal segment is
much concealed; the diminution is due to certain segments at the
extremity being indrawn and serving as a sort of tubular ovipositor in
the female, or curled under the body and altered in form in the other
sex, so as to constitute what is called a "hypopygium." In the female
of Tipulidae the body is terminated by some horny pieces forming an
external ovipositor. In nearly all Diptera the feet are five-jointed; the
claws are well developed, there being placed under each of them a
free pad or membrane, the "pulvillus"; there may be also a median
structure between each pair of claws, of diverse form, the
"empodium."

On the surface of the body of many flies there will be seen an

armature of pointed bristles; these flies are called "chaetophorous";
where no regularly arranged system of such bristles exists the fly is
"eremochaetous." In some families the arrangement of these bristles
is of importance in classification, and a system of description has
been drawn up by Baron Osten Sacken: this branch of descriptive
entomology is known as chaetotaxy.[354]

The wings are of great importance in classifying Diptera; but

unfortunately, like the other parts, they have not received an
exhaustive anatomical study, and Dipterologists are not agreed as to
the names that should be applied to their parts.

Fig. 215—Nervuration of Dipterous wing. A, Wing of a Tipulid,

according to Loew, who uses the following nomenclature: a, costal
nervure; b, mediastinal; c, subcostal; d, radial; e, cubital; f,
discoidal; g, postical; h, anal; i, axillar; x, transverse, y, posterior
transverse, nervure; 1, 2, mediastinal areas; 3, subcostal; 4,
cubital; 5, anterior basal; 6, posterior basal; 7, anal; 8, posterior
marginal; 9, discoidal. B, Wing of an Acalypterate Muscid
(Ortalis), according to Schiner, who uses the following
nomenclature: (nervures, small letters; cells, capital letters): a,
transverse shoulder; b, auxiliary; c to h, first to sixth longitudinal; i,
middle transverse; k, posterior transverse; l, m, n, o, costa; p,
anterior basal transverse; q, posterior basal transverse; r,
rudiment of a fourth nervure; s, axillary incision: A, B, C, first,
second, and third costal cells; D, marginal; E, sub-marginal; F, G,
H, first, second, and third posterior; I, discal; K, L, M, first, second,
and third basal cells; N, anal angle; O, alula.

We give below figures of two systems that have been used by

eminent Dipterologists for the description of the nervures and cells.
The comprehension of these features of the Dipterous wing will be
facilitated by noticing that the wing—being extended at right angles
to the body—is divided by the longitudinal nervures into two great
fields, anterior and posterior, with an interval between them: this
interval is traversed only by a short cross-vein (marked x in Fig. 215
A, and i in B). This cross-vein may be placed near the base or nearer
to the tip of the wing; it is of importance because no nervure in front
of the median area traversed by it can correspond with a nervure
placed behind it in another wing. The very different nature of the
nervuration in the two wings we have figured will readily be
appreciated by an inspection of the parts posterior to the little cross-
vein. On the hind margin of the wing, near the base, there is often a
more or less free lobe (Fig. 215, B, O) called the "alula": still nearer
to the base, or placed on the side of the body, may be seen one or
two other lobes, of which the one nearer the alula is called the
"tegula," or (when a lobe behind it is also present) the "upper tegula,"
(the "antitegula" of Osten Sacken); the other being the "lower
tegula." These two terms are erroneous, the word tegula being
definitely applied to another part of the Insect-body. In speaking of
this structure in the following pages, we have preferred to call it the
"squama."[355] Those Muscidae in which the squama covers the
halter like a hood are called "calypterate." In Fig. 216, we represent
these structures, and in the explanation have mentioned the
synonyms. The terms we think most applicable to the three lobes are
alula, antisquama, squama. The squama may be called "calypter"
when it covers the halter.

Fig. 216—Parts at the base of the wing in Calliphora. a, Anal angle or

lobe of the wing; b, alula; c, antisquama, squama alaris, or
antitegula; d, squama, squama thoracicalis, tegula, calypter, or
calyptron; e, posterior extremity (scutellum) of the mesothorax; f,
scutum of mesothorax.

The halteres—commonly called balancers or poisers—are perhaps

the most characteristic of all the Dipterous structures, though they
are absent in most of the few wingless forms of the Order. Outside
the Diptera similar organs appear to exist only in male Coccidae.
The pair of halteres is placed on the metathorax, one on each of the
pleural regions. They are believed to be the homologues of the hind
wings; Weinland states[356] that certain canals existing in the interior
of the halter correspond to wing-nervures. The halter may be
described as a small rod-like body with a head like a pin, this
terminal part being, however, rather variable in form. We have
already stated that in many Diptera the squama forms a hood, the
position of which leads to the belief that it is an important adjunct to
the halter. Although the exact functions of the halteres are far from
clear, it is certain that they are highly complex bodies, of extremely
delicate structure: they are doubtless sense-organs, possessing as
they do, groups of papillae on the exterior and a chordotonal organ
(a structure for assisting the perception of sound) in the basal part;
each halter is provided with four muscles at the base, and can, like
the wings, execute most rapid vibrations. Seeing that they are the
homologues of wings, it is a remarkable fact that in no Diptera are
they replaced by wings, or by structures intermediate between these
two kinds of organs.

Internal Structure.—Information about the internal anatomy is by no

means extensive. The tracheal system is highly developed, and has
air-sacs connected with it; a large pair at the base of the abdomen
being called aërostats by Dufour. Inside the thoracic spiracles there
are peculiar structures supposed by some to be voice-organs, while
the abdominal spiracles are said to be remarkably simple in
structure. Lowne says that there are ten or eleven pairs of spiracles
in the Blow-fly; one of these, near the base of the wing, is peculiar in
structure, and may not be a true stigma; he calls it a tympanic
spiracle; it seems doubtful whether there are more than seven
abdominal pairs. The alimentary canal is very elongate, and is
provided with a diverticulum, the crop; this is usually called the
sucking stomach, though its function is extremely doubtful. The
Malpighian tubes are four in number, and are very elongate; in
several groups of Nemocera there are, however, five Malpighian
tubes, a number known to occur in only very few other Insects. The
nervous system is remarkable on account of the concentration of
ganglia in the thorax, so as to form a thoracic, in addition to the usual
cephalic, brain. For particulars as to the positions of the ganglia and
the great changes that occur in the lifetime, the student should refer
to Brandt, to Künckel, and to Brauer.[357] Much information as to the
internal anatomy of the Blowfly is given by Lowne, but it is doubtful to
what extent it is applicable to Diptera in general.[358]

Fig. 217—Acephalous larva or maggot of the blow-fly, with the head, a,

extended. (After Lowne.)

The larvae of Diptera are—so far as the unaided eye is concerned—

without exception destitute of any kind of adornment, the vast
majority of them being of the kind known as maggots. None of them
have true thoracic legs; though in the earlier groups, pseudopods or
protuberances of the body that serve as aids in locomotion are
common. Unlike what occurs in other Orders the arrangement of
these pseudopods on the body differs greatly in various forms; in a
few cases they are surmounted by curved hairs. The most important
distinction in external form in Dipterous larvae is that while those that
are thorough maggots possess no visible head, others have a well-
marked one (Fig. 225); these are therefore called "eucephalous":
they have a mouth of the mandibulate type. In some other Dipterous
larvae the head is more or less reduced in size, and in the
acephalous forms there is only a framework of a few chitinous rods
to represent it. The nervous system in the most completely headless
larvae is very remarkable, all the ganglia being concentrated in a
single mass placed in the thorax. The tracheal system exhibits a
great variety; some larvae have stigmata arranged along the sides of
the body after the fashion normal in Insect-larvae; these are called
"peripneustic"; as many as ten pairs of stigmata may be present in
these cases, but nine pairs is much more common. Other larvae
have a pair of stigmata placed at the termination of the body, and
another pair near the anterior extremity, the two pairs communicating
by large tracheal trunks extending the length of the body; these
larvae are said to be "amphipneustic": this is the condition usual in
the more completely acephalous larvae. Others have only the
terminal pair of spiracles, and are styled "metapneustic." Some begin
life in the metapneustic state and afterwards become amphipneustic.
In the aquatic larva of Corethra there are no spiracles, though there
is an imperfect tracheal system. Many Dipterous larvae that live in
water or in conditions that prevent access of air to the body have
remarkable arrangements for keeping the tip of the body in
communication with the atmosphere. The stigmata in metapneustic
and amphipneustic larvae are very remarkable compound structures,
exhibiting however great diversity; their peculiarities and uses are
not well understood; it appears very doubtful whether some of them
have any external opening. Reference may be made, as to the
variety of structure, to Meijere's paper[359] from which we take the
accompanying figure of a posterior stigmatic apparatus in Lipara
lucens. It appears that there is a compound chamber
—"Filzkammer"—terminating externally in lobes or fingers
—"Knospen" and appearing as marks on the outer surface: this
chamber is seated on a tracheal tube, and is, Meijere thinks,
probably a secondary growth of the trachea coming to the outer
surface. It is traversed by what may be considered the original
tracheal tube, opening externally as an external stigmatic scar
—"Stigmennarbe"—and with a second or inner scar placed
internally. We may conclude from what is already known that these
structures will be found to differ in the same larva according to the
stage of its development.

Fig. 218—The posterior stigma of the larva of Lipara lucens. a, One of

the three "Knospen" or lobes; b, external stigmatic scar; c, internal
scar; d, stigmatic chamber (Filzkammer); e, trachea. (After
Meijere.)
An extremely valuable summary of the characters and variety of
Dipterous larvae has been given by Brauer,[360] from which it
appears that the larvae of the first half of the family exhibit great
variety and have been much studied, while the more purely maggot-
like forms of the Muscidae have, with one or two exceptions, been
little investigated.

The pupal instar is of two distinct kinds. First, we meet with a pupa
like that of Lepidoptera, viz. a mummy-like object, or pupa obtecta, in
which there is a crisp outer shell, formed in part by the adherent
cases of the appendages of the future imago. This condition, with a
few exceptions to be subsequently noticed, obtains in the Nemocera
and Brachycera. It is exhibited in various degrees of perfection,
being most complete in Tipulidae; in other forms the shell is softer
and the appendages more protuberant. The second kind of pupa is
found in the Cyclorrhaphous flies; it has externally no marks except
some faint circular rings and, frequently, a pair of projections from
near one extremity of the body; occasionally there is a single
prominence at the other extremity of the body. This condition is due
to the fact that the larva does not escape from the skin at the last
ecdysis, but merely shrinks within it, so that the larval skin, itself
contracted and altered by an excretion of chitin, remains and forms a
perfect protection to the included organism. This kind of pupa looks
like a seed, and is well exemplified by the common Blow-fly. The
capacity for entering on such a condition is evidently correlative with
the absence of a larval head. The metamorphosis in this curious little
barrel goes on in a different manner to what it does in the pupa
obtecta. A good name for the whole structure of this instar has not
been found. Older authors called it "pupa coarctata," or "nympha
inclusa"; Brauer speaks of it as a "compound pupa"; ordinarily in our
language it is called a "puparium," a term which is more applicable to
the case alone.

In species having a pupa obtecta the larval skin is cast after the chief
processes of the external metamorphosis have occurred, and then
an exudation of chitin hardens the general surface. In the
"compound pupa" of the Blow-fly there is for a considerable period
no formed pupa at all, but merely a shell or case containing the
results of histolysis and the centres for regeneration of new organs;
the chitin-exudation to the exterior of the larval skin occurs in the
early part of the series of metamorphic changes, and the organism
breaks down to a cream within the shell thus formed, and then
gradually assumes therein the condition of a soft, nymphoid pupa.
The exceptional conditions previously referred to as exhibited by a
few forms are certain cases in which a more or less perfect pupa
obtecta is found within the last larval skin, as is the case in
Stratiomys. Another highly remarkable condition exists in the
Hessian fly, and a few other Cecidomyiids, where the Insect
apparently makes an exudation which it uses as a covering case,
independent of the larval skin; this latter being subsequently shed
inside the case, so that this condition of coarctate pupa differs from
that we have described as existing in Cyclorrhaphous flies, although
the two are superficially similar. In the Pupipara the larval stage is
passed in the body of the mother, which produces a succession of
young, nourished one at a time by the secretion of glands; this young
is born as a full-grown larva that becomes at once a pupa.

Metamorphosis.—As it is in Diptera that the phenomena of Insect-

metamorphosis have reached their highest development we
endeavoured to give some idea of their nature in the previous
volume, therefore we need give only a brief sketch of the chief
features of Dipterous metamorphosis. The Blow-fly undergoes a
rapid embryonic development, the later stages of which are, on the
whole, of a retrogressive nature. On the emergence of the young
maggot it feeds up rapidly, the rapidity varying greatly according to
circumstances, and then when full-grown rests. While resting, a
process of internal liquefaction, called histolysis, is going on, and the
maggot contracts and exudes an excretion that hardens its skin. At
the time this hard skin has become complete, or soon after, the
maggot inside has dissolved into a cream contained in a sac inside
the shell; this cream becomes reconstituted into a fly by a gradual
process of growth and development of certain minute portions of the
body—the imaginal discs or folds, the histoblasts and neuroblasts
that were exempt from the histolytic process: in the early stages of
the reconstitution the general structure is, of course, altogether
vague, and this condition—purely one of transition—is called the
pronymph; the nymph becomes gradually developed: it corresponds
vaguely with the pupa obtecta of the early groups of Diptera, but is
soft like the pupa of Hymenoptera. This nymph gradually develops
into the fly itself, the external parts being first completed and the
internal organs elaborated subsequently. The sexual organs do not
undergo metamorphosis like other internal organs, there being a
gradual (though irregular or interrupted) growth of them in the young
larva, till they are completed some time after the emergence of the
perfect fly. The processes in the Blow-fly have been studied by
numerous able histologists of various nationalities, and have recently
been described by Lowne in our own language.[361] Comparatively
little has been done in studying the corresponding phenomena in
other Diptera. Weismann has investigated the development of
Corethra, and Miall that of Chironomus. These two flies belong to a
division of Diptera different from that which includes the Blow-fly, and
they display a condition of the metamorphic processes allied to what
occurs in Lepidoptera, as well as to that which takes place in the
Blow-fly. Imaginal folds are formed, but they only appear much later
in the life, and they are much less distant from the positions they will,
when developed, occupy in the imago. In Chironomus, according to
Miall, the imaginal folds only appear in the last larval instar, but they
grow with such rapidity that the legs and wings of the future fly can
be distinguished in the larva, even before pupation; thus when the
activity of the larva ceases but little change is required to complete
the obtected pupa. In the Blow-fly some of the imaginal folds have
been traced back to the embryo; how many centres for the new
growth there may be is uncertain, for though there are upwards of
sixty for the outer body, the number of regenerative centres for the
internal organs is not ascertained. The peculiar central nervous
mass, mentioned in our remarks on the larva, consists of two kinds
of tissue mixed together in a complex manner; one of these kinds is
functionally active during the larval life and at the metamorphosis
undergoes histolysis, while the other, or embryonic, portion develops
into the nervous system of the fly.
It forms no part of our task to deal with general subjects, but we may
be pardoned for calling attention to the bearing the metamorphosis
of the higher Diptera has on our ideas of heredity in Insects. The fly
bears no resemblance whatever to the larva, and is only obtained by
the organic destruction of the latter, which occurs before the
perfection of the sexual organs takes place, and yet the fly
reproduces itself only secondarily, but primarily gives rise to the
totally different larva. It is supposed that the larval structures have
been gradually acquired, and yet they are transmitted with the
utmost faithfulness by the totally different fly. We can only conclude
that that which is bequeathed in each species is the early state of a
particular process of development from which the subsequent stages
follow necessarily if the developing organism be placed in conditions
having on it influences like to those that influenced the ancestors.

Classification.—The classification of Diptera is as yet very

imperfect. Formerly they were divided into two great groups,
Nemocera and Brachycera, according to the structure of the
antennae, as previously mentioned. This division has been
abandoned, and the term Brachycera is now applied to only a small
part of the old section that bore the name. The primary division
usually adopted at present is into Orthorrhapha and Cyclorrhapha.
The characters of these two groups are based on the nature of the
metamorphosis, and have been gradually elaborated by Brauer in
various memoirs.[362] The Orthorrhapha includes the forms with
obtected pupae, the Cyclorrhapha those with a nymph-compound,
as previously described. This distinction is of great importance, but
unfortunately it is difficult to apply to the fly itself; the only character
that can be used in connection with the imago is the existence of a
suture over the insertion of the antennae in a portion, but not all, of
the Cyclorrhapha.[363] The next set of divisions used by Brauer
divides the Order into four sections, viz. 1. Orthorrhapha
Nematocera, 2. O. Brachycera, 3. Cyclorrhapha Aschiza, 4. C.
Schizophora. As these four groups are recognised more readily than
the two major groups the student will do well at first to disregard the
primary division and consider the Diptera as divisible into four great