by Hampson in the sub-family Phaudinae of Zygaenidae characterised by the
absence of the mouth-parts. The Himantopteridae are small moths, and have
the scales on the wings very imperfect and hair-like; the hind wings form
long slender tails, so that the Insects scarcely look like moths. They are
peculiar to India and Africa. In the South African genus _Dianeura_
(belonging really to Phaudinae) also the wings are scaleless and nearly
transparent.
FAM. 18. HETEROGYNIDAE.—Consists of the single genus _Heterogynis_ which
has hitherto been found only in the south of Europe. This is an important
form connecting Zygaenidae and Psychidae. The larvae resemble those of
Zygaena, and construct an oval cocoon for their metamorphosis. The male
issues as a small moth of smoky colour, the scales being but imperfect; the
female chrysalis shows no trace of any appendages, and the imago is
practically a maggot, and never leaves the cocoon; in it she deposits her
eggs, and the young larvae hatch there.[275]
FAM. 19. PSYCHIDAE.—Small, or moderate-sized moths, with imperfect scales,
and little or no colour beyond certain shades of duskiness; the sexes very
different, the female being wingless and sometimes quite maggot-like; the
male often with remarkable, bipectinate antennae, the branches sometimes
very long and flexible. Larva inhabiting a case that it carries about. This
family consists of Insects unattractive in appearance but presenting some
points of great interest. It is frequently stated that the Psychidae are
destitute of scales, but Heylaerts states[276] that, in addition to hairs,
scales of a more or less imperfect formation are present in all, but that
they are, like those of some Sphingidae (_Macroglossa_), very easily
detached. There is much difference in the females, some having
well-developed legs, while others are not only apterous, but are bare and
destitute of appendages like a maggot, while in certain cases (Fig. 196,
G), the head is reduced in size and is of peculiar form so as to make the
Insect look really like the larva of one of the parasitic Diptera. These
females never leave their cases, but deposit their eggs {393}therein, and
inside, also, their former pupa-skin; and here the young hatch; the
peculiar little larvae are very numerous, and it is suggested that they
make a first meal on the body of their parent, but this we believe has not
been satisfactorily ascertained. Great differences as to the condition of
the legs, antennae, etc., are said to exist in species of the same genus.
There is also a remarkable diversity in the pupae of the females; the male
sex being normal in this respect. Some of the female pupae are destitute of
wing-sheaths and all other appendages, while others are said to possess
them, though there are no wings at all in the imago (_Fumea_, _e.g._).[277]
Great difficulties attend the study of these case-bearing Insects, and
several points require careful reconsideration, amongst them the one we
have just mentioned. The males fly rapidly in a wild manner, and may
sometimes be met with in swarms; their lives are believed to be very brief,
rarely exceeding a couple of days, and sometimes being limited to a few
hours.
[Illustration: Fig. 196—Metamorphosis of _Monda rhabdophora_. Ceylon. A,
Larva in case, nat. size; B, larva itself, magnified; C, case of female
during pupation; D, case of male during pupation; E, female pupa,
magnified; F, male moth, nat. size; G, female moth, magnified. (From
unpublished drawings by Mr. E. E. Green).]
The larvae are called basket-worms, and their baskets or cases are well
worthy of attention. Their variety is remarkable; the most extraordinary
are some of the genus _Apterona_ Fig. 197, B, which perfectly resemble the
shells of Molluscs such as snails; indeed, the specimens in the collection
at the British Museum were sent there as shells. This case is not, like
those of other Psychidae, constructed of earth or vegetable matter, but is
of silk and is in texture and appearance exactly {394}like the surface of a
shell. _Psyche helix_ is, according to Ingenitzky,[278] found in great
numbers near Lake Issyk-kul in Central Asia, where the larvae feed, in
their snail-shell-like cases, on a grass, just like snails. Only females
could be reared from these larvae. The case of _Chalia hockingi_ (Fig. 197,
C) consists of little pieces of wood cut to the proper lengths, and
spirally arranged, so as to form a construction that would be quite a
credit to our own species. In some of the Canephorinae we meet with long
cylindrical cases, like those of Caddis-worms, or of Tineid larvae.
[Illustration: Fig. 197—Baskets, or cases, of Psychidae. A, _Amicta
quadrangularis_; B, _Apterona_ (or _Cochlophora_) _valvata_; C, _Chalia
hockingi_.]
Riley has given an account of several points in the structure and natural
history of one of the North American basket- or bag-worms, _Thyridopteryx
ephemeraeformis_; one of his points being the manner in which the newly
hatched larva forms its case.[279] This question has also been discussed by
Packard.[280] The larvae when hatched in unnatural conditions will make use
of fragments of paper, cork, etc., for the case; the act of construction
takes one or two hours, and the larva does not eat till the case is
completed. It walks in a peculiar manner, the legs of the third pair being
moved forwards together, as if they were the prongs of a fork.
This family is already one of considerable extent, but its study, as
already remarked, is but little advanced. Some naturalists are inclined to
place it among the Tineidae, but it is connected with Zygaenidae by means
of Heterogynidae. Mr. Meyrick divides it, placing _Psyche_ and
_Sterrhopteryx_ (the forms representing, according to his ideas, the family
Psychidae in Britain) in the series Psychina which includes Zygaenidae. He
{395}removes the other British genera, _Fumea_, and _Epichnopteryx_, to
Tineidae near _Solenobia_ and _Taleporia_. The group Canephorinae, to which
the two genera in question belong, was long since separated from Psychidae
by Herrich-Schäffer, but this course was condemned by Heylaerts.
Parthenogenesis has been thought by some to occur in numerous species in
this family, but Heylaerts says that it is limited to _Apterona crenulella_
var. _helix_, and even of this species males are found in certain
localities.
FAM. 20. COSSIDAE (_Goat-Moths_, or _Carpenter-Worms_).—Moths of moderate,
or rather large size, without proboscis, frequently with a dense covering
of matted, imperfect scales; the pattern being vague. The larvae bore into
trees in which they often make large burrows, leaving holes from which sap
exudes. Our common Goat-moth is a good specimen of this family, which is a
very widely distributed one. The Australian genus _Ptilomacra_ has very
large, pectinated antennae in the male. The larvae of Cossidae are nearly
bare of clothing and are unadorned; they form a slight cocoon of silk mixed
with gnawed wood. The pupa of the Goat-moth is remarkable for the great
development of the rows of teeth on the dorsal aspects of the segments of
the abdomen, and for the absence of consolidation in this part, six of the
intersegmental incisions being free, and the ventral aspect almost
membranous. Very little is known as to other pupae of the family. It is
believed that the generations of these Insects are fewer than usual, the
growth of the larva occupying a period of two or three years. The larva of
_Zeuzera aesculi_ forms a temporary cocoon in which it passes a
winter-sleep, before again feeding in the spring.[281] It is a moot
question whether the Zeuzeridae should be separated from the Cossidae or
not. The group includes our Wood-leopard moth, which, like many other
Zeuzerids, is spotted in a very striking but inartistic manner. The
position the family Cossidae should occupy in an arrangement of the
Lepidoptera is a very difficult question. Some consider the Insects to be
allied to Tortricidae. The wing-nervuration of _Cossus_ is very peculiar
and complex, there being four or five cells on the front wing, and three on
the hind one. Meyrick places Zeuzeridae as a family of his series Psychina,
but separates Cossidae proper (he calls them Trypanidae) as a family of the
series Tortricina.
{396}FAM. 21. ARBELIDAE.—Closely allied to Cossidae, but without frenulum,
and with less complex wing-nervures. A small family believed to be similar
to Cossidae in the life-history. The tropical African Arbelidae are
considered by Karsch to be a distinct family, Hollandiidae.
FAM. 22. CHRYSOPOLOMIDAE.—This family has been established by Dr.
Aurivillius[282] for an African genus, allied in wing-nervuration to
Cossidae; the Insects are like Lasiocampidae.
FAM. 23. HEPIALIDAE (_Ghost-_ and _Swift-Moths_).—Moths of very diverse
size, some gigantic; wings not fitting together well at the bases; without
a frenulum; no proboscis; the scales imperfect; the nervures complex. The
Hepialidae are extremely isolated amongst the Lepidoptera; indeed, they
have really no allies; the conclusion that they are connected with the
Micropterygidae being certainly erroneous. Although but small in
numbers—only about 150 species being known—they exhibit a remarkable
variety in size and colour. Many are small obscure moths, while others are
of gigantic size—six or seven inches across the wings—and are amongst the
most remarkably coloured of existing Insects. The great _Charagia_ of
Australia, with colours of green and rose, bearing white spots, are
remarkable. The South African _Leto venus_ is of large size, and has an
astonishing supply of glittering metallic splashes on the wings, making a
barbaric but effective display. The South Australian _Zelotypia staceyi_,
of enormous size, is also a handsome moth; but the majority of species of
the family are adorned only in the feeblest manner.
Very little is known as to the larvae; they are either subterranean,
feeding on roots, or they live in the wood of trees and shrubs. They are
nearly bare, and are apparently the lowest type of Bombycid larva. At the
same time, it would appear there is considerable variety amongst them.
Packard says[283] the young larva of _Hepialus mustelinus_ has the
arrangement of setae that is normal in Tineidae. The larva of _H. humuli_
seems to be a very simple form, but _H. hectus_ shows a considerable amount
of divergence from it. They probably live for several years; the larva of
_H. argenteo-maculatus_ in North America lives for three years, at first
eating the roots of Alder and then entering the {397}stems. The pupae are
also peculiar. They are of unusually elongate, cylindrical form, with
comparatively feeble integument, but with a considerable development of
chitinous, elevated, toothed ridges, on the dorsal aspect, and a very
strong ridge of this kind on the ventral surface of the seventh segment;
the wing-sheaths are short; it is very difficult to distinguish the full
number of abdominal segments. These pupae are remarkably agile, and by
wriggling and kicking are able to move a considerable distance; it is said
that they can force themselves to the surface even when the superficial
soil is quite hard. We cannot consider this pupa naturally placed amongst
either the pupae obtectae or incompletae of Chapman.
[Illustration: Fig. 198—Pupa of _Hepialus lupulinus_. Britain. A, Ventral;
B, dorsal aspect.]
We have already remarked that little is known as to the life-histories. The
species are probably prolific, a female of _H. thule_ having been known to
deposit more than 2000 eggs. Of the Australian forms little more is
known[284] than that they live in the wood of trees and shrubs, and are
rapidly disappearing; we may fear that some are extinct without ever having
been discovered, and others, also unknown but still existing, may disappear
only too soon; the wasteful destruction of timber in Australia having been
deplorable.
The peculiar habits of the Hepialidae are not likely to bring the Insects
to the net of the ordinary collector, and we believe they never fly to
light, hence it is probable that we are acquainted with only a small
portion of the existing species; their distribution is very wide, but
Australia seems to be their metropolis, and in New Zealand twelve species
are known. The genera as at present accepted are remarkable for their wide
distribution. _Leto_ is said to occur in South Africa and in the {398}Fiji
Islands; but we must repeat that the study of these interesting Insects is
in a very primitive state, and our present knowledge of their distribution
may be somewhat misleading.
The habits of the European _Hepialus_ in courtship have been observed to a
considerable extent and are of great interest, an astonishing variety and a
profound distinction in the methods by which the sexes are brought together
having been revealed.
_H. humuli_, our Ghost-moth, is the most peculiar. Its habits were detected
by Dr. Chapman.[285] The male is an Insect of exceptional colour, being
white above, in consequence of a dense formation of imperfect scales; the
female is of the brownish tints usual in Swift-moths. In the month of June
the male selects a spot where he is conspicuous, and hovers persistently
there for a period of about twenty minutes in the twilight; his colour has
a silvery-white, glistening appearance, so that the Insect is really
conspicuous notwithstanding the advanced hour. Females may be detected
hovering in a somewhat similar manner, but are not conspicuous like the
male, their colour being obscure; while so hovering they are ovipositing,
dropping the eggs amongst the grass. Females that have not been fertilised
move very differently and dash about in an erratic manner till they see a
male; they apparently have no better means of informing the hovering male
of their presence than by buzzing near, or colliding with him. Immediately
this is done, the male abandons his hovering, and coupling occurs. There
can be little doubt that the colour of the male attracts the female; but
there is a variety, _hethlandica_, of the former sex coloured much like the
female, and in some localities varieties of this sort are very prevalent,
though in others the species is quite constant. This variation in the
colour of the males is very great in Shetland,[286] some being quite like
the females. In _H. hectus_ the two sexes are inconspicuously and similarly
coloured. The male hovers in the afternoon or evening in a protected spot,
and while doing so diffuses an agreeable odour—said by Barrett to be like
pine-apple—and this brings the female to him, much in the same manner as
the colour of _H. humuli_ brings its female. The hind legs of the male
{399}are swollen, being filled with glands for secreting the odorous
matter.[287] This structure has led to the suggestion of the generic name
_Phymatopus_ for the Insect. Turning to other species of the genus, we find
that the normal relative rôles of the sexes are exhibited, but with
considerable diversity in the species. In _H. lupulinus_ the males fly
about with rapidity, while the female sits on a stem and vibrates her
wings; she thus attracts the males, but they do not perceive her unless
happening to come within three or four feet, when they become aware of her
proximity, search for and find her. It is doubtful whether the attraction
is in this case the result of an odour; it would appear more probable that
it may be sound, or that the vibration of the wings may be felt by the
male.
In _H. sylvinus_, _H. velleda_ and _H. pyrenaicus_ less abnormal modes of
attracting the males occur, the individuals of this latter sex assembling
in great numbers at a spot where there is a female. In the first of the
three species mentioned the female sits in the twilight on the stem of some
plant and vibrates the wings with rapidity; she does not fly; indeed,
according to Mr. Robson, she does not till after fertilisation move from
the spot where she emerged. In _H. pyrenaicus_ the female is quite
apterous, but is very attractive to the males, which as we have said,
assemble in large numbers near her. Thus within the limits of these few
allied forms we find radically different relations of the sexes.
1. The male attracts the female—(A) by sight (_H. humuli_); (B) by odour
(_H. hectus_).
2. The female attracts the male—(A) by vibration of wings (_H. lupulinus_
and _H. sylvinus_); (B) without vibration, but by some means acting at a
distance (_H. velleda_, _H. pyrenaicus_).
Little or nothing is known as to the habits of the great majority of the
more remarkable forms of the family. The gigantic Australian forms are
believed to be scarcely ever seen on the wing.
The Hepialidae differ from other Lepidoptera by very important anatomical
characters. The absence of most of the {400}mouth-parts is a character
common to them and several other divisions of Lepidoptera; but the labial
palpi are peculiarly formed in this family, being short and the greater
portion of their length consisting of an undivided base, which probably
represents some part of the labium that is membranous in normal
Lepidoptera. The thoracic segments are remarkably simple, the three
differing less from one another than usual, and both meso- and meta-notum
being much less infolded and co-ordinated. The wings are remarkable for the
similarity of the nervuration of the front and hind wings, and by the cell
being divided by longitudinal nervules so as to form three or four cells.
On the inner margin of the front wing there is near the base an incision
marking off a small prominent lobe, the jugum of Prof. Comstock. Brandt
mentions the following anatomical peculiarities,[288] viz. the anterior
part of the alimentary canal is comparatively simple; the respiratory
system is in some points like that of the larva; the heart is composed of
eight chambers; the appendicular glands of the female genitalia are
wanting. The testes remain separate organs throughout life. The chain of
nerve ganglia consists of the supra- and infra-oesophageal, three thoracic,
and five abdominal, ganglia, while other Lepidoptera have four abdominal.
FAM. 24. CALLIDULIDAE.—A small family of light-bodied diurnal moths having
a great resemblance to butterflies. In some the frenulum is present in a
very rudimentary condition, and in others it is apparently absent.
_Cleosiris_ and _Pterodecta_ are very like butterflies of the Lycaenid
genus _Thecla_. Although fifty species and seven or eight genera are known,
we are quite ignorant of the metamorphoses. Most of the species are found
in the islands of the Malay Archipelago, but there are a few in East India.
FAM. 25. DREPANIDAE (or DREPANULIDAE). (_Hook-tips_).—The larger moths of
this family are of moderate size; many of the species have the apex of the
front wing pointed or even hooked; some have very much the appearance of
Geometrid moths; they resemble very different members of that family.
_Oreta hyalodisca_ is remarkable on account of the very large, transparent
patch on each front wing, though the other species of the genus have
nothing of the sort. In the genus _Deroca_ we {401}find Insects with the
scales imperfect, they being few and small and approximating in form to
hairs; in _D. hyalina_ scales are nearly entirely absent. In other genera,
_e.g._ _Peridrepana_, _Streptoperas_, there is only a very inferior state
of scale-formation. The few larvae that are known are peculiar; they are
nearly bare of hair, without the pair of terminal claspers, while the body
is terminated by a long tubular process. They form a slight cocoon among
leaves.
The members of the family were formerly much misunderstood, and were
assigned to various positions in the Order. There are now about 30 genera,
and 150 species known, the geographical distribution of the family being
very wide. In Britain we have half a dozen species. _Cilix glaucata_
(better known as _C. spinula_) is said "to undoubtedly imitate" the
excrement of birds. No doubt the Insect resembles that substance so as to
be readily mistaken for it. This Insect has a very wide distribution in
North America, Europe and East India, and is said to vary so much in the
structure of its organs as to justify us in saying that the one species
belongs to two or three genera.
[Illustration: Fig. 199—Mature larva of _Apoda testudo_, on beech-leaf.
Britain.]
FAM. 26. LIMACODIDAE (or EUCLEIDAE).—These are somewhat small moths, of
stout formation, sometimes very short in the body, and with rather small
wing-area. The family includes however at present many Insects of diverse
appearance; there are numerous forms in which apple-green is a prominent
colour; some bear a certain resemblance to the Swifts, others to Noctuids;
some, _Rosema_ and _Staetherinia_, are of extraordinary shapes; certain
very small forms, _Gavara_, _Ceratonema_, resemble Tortricids or Tineids; a
few even remind one of Insects of other Orders; so that the group is a
mimetic one. _Nagoda nigricans_ (Ceylon) has the male somewhat like a
Psychid, while the female has a different system of coloration and
wing-form. In _Scopelodes_ the palpi are in both sexes remarkable;
elongated, stiff, directed upwards and brush-like at the tip. Altogether
there are about 100 genera and 400 species known; the distribution of the
family is very wide {402}in both hemispheres, but these Insects do not
occur in insular faunas. In Britain we have two genera, _Heterogenea_ and
_Apoda_ (better known as _Limacodes_[289]), each with a single species.
[Illustration: Fig. 200—Larva of _Apoda testudo_ just hatched. A, Dorsal
view of larva; B, C, D, a spine in different states of evagination. All
magnified. (After Chapman.)]
The early stages of these Insects are of great interest. The eggs, so far
as known, are peculiar flat oval scales, of irregular outline and
transparent; we have figured an example in Vol. V. Fig. 83. The eggs of the
same moth are said to vary much in size, though the larvae that emerge from
them differ little from one another in this respect. The latter are
peculiar, inasmuch as they have no abdominal feet, and the thoracic legs
are but small; hence the caterpillars move in an imperceptible gliding
manner that has suggested for some of them the name of slug-worms. The
metamorphoses of a few are known. They may be arranged in two groups; one
in which the larva is spinose or armed with a series of projections and
appendages persisting throughout life; while in the members of the second
group the spines have only a temporary existence. At the moment the young
larva of _Apoda testudo_ emerges from the egg it has no conspicuous spines
or processes, and is an extremely soft, colourless creature,[290] but it
almost immediately displays a remarkable system of complex spines. These
really exist in the larva when it is hatched, and are thrust out from pits,
as explained by Dr. Chapman. In the succeeding stages, the spines become
modified in form, and the colour of the body and the nature of {403}the
integument are much changed, so that in the adult larva (Fig. 199) the
spines have subsided into the condition of mere prominences, different in
colour from the rest of the surface. These larvae appear to be destitute of
a head, but there really exists a large one which is retracted, except
during feeding, into the body; the five pairs of abdominal feet of the
larvae of allied families are replaced by sucker-like structures on the
first eight abdominal segments. The spinneret of the mouth is not a pointed
tubular organ, but is fish-tailed in shape, and hence disposes the silky
matter, that aids the larva in moving on the leaves, in the form of a
ribbon instead of that of a thread. It has been stated that these peculiar
larvae "imitate" the coloured galls frequently found on the leaves of
trees. The North American forms of this family have very varied and most
extraordinary larvae.[291] In the pretty and conspicuous larva of _Empretia
stimulea_, the tubercles or processes of the body are, in the later stages,
armed with hairs, that contain a poisonous or irritating fluid, said to be
secreted by glands at the bases of the processes. These hairs are readily
detached and enter the skin of persons handling the caterpillars. The larva
of the North American Hag-moth, _Phobetron pithecium_, is a curious object,
bearing long, fleshy appendages covered with down. Hubbard makes the
following statement as to the instincts of this larva:[292]—"The hag-moth
larvae do not seek to hide away their cocoons, but attach them to leaves
and twigs fully exposed to view, with, however, such artful management as
to surroundings and harmonising colours that they are of all the group the
most difficult to discover. A device to which this Insect frequently
resorts exhibits the extreme of instinctive sagacity. If the caterpillar
cannot find at hand a suitable place in which to weave its cocoon, it
frequently makes for itself more satisfactory surroundings by killing the
leaves, upon which, after they have become dry and brown in colour, it
places its cocoon. Several of these caterpillars unite together, and
selecting a long and vigorous immature shoot or leader of the orange tree,
they kill it by cutting into its base until it wilts and bends over. The
leaves of a young shoot in drying turn a light tan-color, which
{404}harmonises most perfectly with the hairy locks of the caterpillar
covering the cocoon. The latter is, consequently, not easily detected, even
when placed upon the exposed and upturned surface of the leaf."
The cocoons of Limacodidae are unusually elaborate, the larva forming a
perfect lid in order to permit itself to escape when a moth. Chapman states
that the larva lies unchanged in the cocoon all winter, moulting to a pupa
in the spring, and that the pupa escapes from the cocoon previous to the
emergence of the moth.[293] Both Chapman and Packard look on the family as
really nearer to Microlepidoptera than to Bombyces; Meyrick (calling it
Heterogeneidae) places it at the end of his series Psychina next
Zygaenidae.
We may allude here to the little moths, described by Westwood under the
name of _Epipyrops_,[294] that have the extraordinary habit of living on
the bodies of live Homopterous Insects of the family Fulgoridae in India.
What their nutriment may be is not known. The larva exudes a white
flocculent matter, which becomes a considerable mass, in the midst of which
the caterpillar changes to a pupa. Westwood placed the Insect in Arctiidae;
Sir George Hampson suggests it may be a Limacodid, and this appears
probable.
FAM. 27. MEGALOPYGIDAE (or LAGOIDAE).—The American genera, _Megalopyge_ and
_Lagoa_, are treated by Berg and by Packard[295] as a distinct family
intermediate between Saturniidae and Limacodidae. The larva is said by the
latter authority to have seven pairs of abdominal feet instead of five
pairs—the usual number in Lepidoptera. When young the caterpillars of
_Lagoa opercularis_ are white and resemble a flock of cotton wool. When
full grown the larva presents the singular appearance of a lock of hair,
moving in a gliding, slug-like manner. Under the long silky hair there are
short, stiff, poison-hairs. The larva forms a cocoon, fitted with a hinged
trap-door for the escape of the future moth. This curious larva is
destroyed by both Dipterous and Hymenopterous parasites.
FAM. 28. THYRIDIDAE.—A small family of Pyraloid moths, exhibiting
considerable variety of form and colour, frequently with hyaline patches on
the wings. They are mostly small {405}Insects, and contain no very striking
forms. Some of them look like Geometrids of various groups. The family is
widely distributed in the tropical zone, and includes 25 genera, of which
_Rhodoneura_, with upwards of 100 species, is the chief one. The larvae are
said to be similar to those of Pyralidae. This family is considered by
Hampson and Meyrick to be ancestral to butterflies.[296]
[Illustration: Fig. 201.—Lappet-moth, _Gastropacha quercifolia_, ♀.
Britain.]
FAM. 29. LASIOCAMPIDAE (_Eggers_, _Lappet-moths_). Usually large Insects
densely covered with scales, without frenulum, but with the costal area of
the hind wing largely developed, and the male antennae beautifully
pectinate, Lasiocampids are easily recognised. They are well known in
Britain, though we have but few species. The flight of some of the species
is powerful, but ill-directed, and the males especially, dash about as if
their flight were quite undirected; as indeed it probably is. The
difference in the flight of the two sexes is great in some species. In the
genus _Suana_ and its allies we meet with moths in which the difference in
size of the two sexes is extreme; the males may be but 1½ inches across the
wings, while the very heavy females may have three times as great an
expanse. Kirby separates these Insects to form the family Pinaridae; it
includes the Madagascar silkworm, _Borocera madagascariensis_. The African
genus _Hilbrides_ is remarkable for the wings being destitute of scales,
and consequently transparent, and for being of very slender form like a
butterfly. The eggs of Lasiocampidae are smooth, in certain cases spotted
in an irregular manner like birds' eggs. Sometimes the parent covers them
with hair. The larvae are clothed with a soft, woolly hair, as well as with
a shorter and stiffer kind, neither beautifully arranged nor highly
coloured, and thus differing from the caterpillars of Lymantriidae; this
hair in some cases has very irritating properties. Cocoons of a close and
compact nature are formed, and hairs from the body are frequently mixed
with the cocoon. In {406}some species the walls of the cocoons have a firm
appearance, looking very like egg-shell—a fact which is supposed to have
given rise to the name of Eggers. Professors Poulton and Meldola have
informed us that this appearance is produced by spreading calcium oxalate
on a slight framework of silk, the substance in question being a product of
the Malpighian tubes.[297] In various families of Lepidoptera it happens
that occasionally the pupa exists longer than usual before the appearance
of the perfect Insect, and in certain members of this family—notoriously in
_Poecilocampa populi_, the December moth—this interval may be prolonged for
several years. There is not at present any explanation of this fact. It may
be of interest to mention the following case:—From a batch of about 100
eggs deposited by one moth, in the year 1891 (the Puss-Moth of the family
Notodontidae), some sixty or seventy cocoons were obtained, the feeding up
of all the larvae having been effected within fourteen days of one another;
fourteen of the Insects emerged as moths in 1892; about the same number in
1893; in 1894, twenty-five; and in 1895, eleven emerged. Lasiocampidae is a
large family, consisting of some 100 genera and 500 or more species, and is
widely distributed. It is unfortunately styled Bombycidae by some
naturalists.
FAM. 30. ENDROMIDAE.—The "Kentish glory," _Endromis versicolor_, forms this
family; it is a large and strong moth, and flies wildly in the daytime in
birch-woods. The larva has but few hairs, and is said when young to assume
a peculiar position, similar to that of saw-fly larvae, by bending the head
and thorax backwards over the rest of the body.
FAM. 31. PTEROTHYSANIDAE.—Consists of the curious East Indian genus
_Pterothysanus_, in which the inner margins of the hind wings are fringed
with long hairs. They are moths of slender build, with large wing-expanse,
black and white in colour, like Geometrids. There is no frenulum.
Metamorphoses unknown.
FAM. 32. LYMANTRIIDAE.—(Better known as LIPARIDAE). These are mostly small
or moderate-sized moths, without brilliant colours; white, black, grey and
brown being predominant; with highly-developed, pectinated antennae in the
male. The larva is very hairy, and usually bears tufts or brushes of
shorter hairs, {407}together with others much longer and softer, these
being sometimes also amalgamated to form pencils; the coloration of these
larvae is in many cases very conspicuous, the tufts and pencils being of
vivid and strongly contrasted colours. Some of these hairy larvae are
poisonous. A cocoon, in which much hair is mixed, is formed. The pupae are
remarkable, inasmuch as they too are frequently hairy, a very unusual
condition in Lepidoptera. The Lymantriidae is one of the largest families
of the old group Bombyces; it includes some 180 genera and 800 species, and
is largely represented in Australia. _Dasychira rossii_ is found in the
Arctic regions. In Britain we have eight genera represented by eleven
species; the Gold-tails, Brown-tails and Vapourer-moths being our commonest
Bombyces, and the latter being specially fond of the London squares and
gardens, where its beautiful larva may be observed on the leaves of roses.
Most of the Lymantriidae are nocturnal, but the male Vapourer-moth flies in
the daytime. In this family there are various species whose females have
the wings small and unfit for flight, the Insects being very sluggish, and
their bodies very heavy. This is the state of the female of the
Vapourer-moth. The males in these cases are generally remarkably active,
and very rapid on the wing.
Some of these moths increase in numbers to an enormous extent, and commit
great ravages. _Psilura monacha_—the Nun, "die Nonne" of the
Germans,[298]—is one of the principal troubles of the conservators of
forests in Germany, and great sums of money are expended in combating it;
all sorts of means for repressing it, including its infection by fungi,
have been tried in vain. The caterpillars are, however, very subject to a
fungoid disease, communicated by natural means. It is believed, too, that
its continuance in any locality is checked after a time by a change in the
ratio of the two sexes. It is not a prolific moth, for it lays only about
100 eggs, but it has been shown that after making allowance for the
numerous individuals destroyed by various enemies, the produce of one moth
amounts in five generations to between four and five million individuals.
The larva feeds on Coniferae, and on many leafy trees and shrubs. The young
{408}larva is provided with two sets of setae, one set consisting of very
long hairs, the other of setae radiating from warts; each one of this
second set of spines has a small bladder in the middle, and it has been
suggested that these assist in the dissemination of the young caterpillars
by atmospheric means.[299] These aerostatic setae exist only in the young
larva. The markings of the moth are very variable; melanism is very common
both in the larva and imago; it has been shown conclusively that these
variations are not connected, as black larvae do not give a larger
proportion of black moths than light-coloured caterpillars do. In England
this moth is never injurious. A closely allied form, _Ocneria dispar_, was
introduced by an accident into North America from Europe about thirty years
ago; for twenty years after its introduction it did no harm, and attracted
but little attention; it has, however, now increased so much in certain
districts that large sums of money have been expended in attempting its
extirpation.
_Dasychira pudibunda_ has occasionally increased locally to an enormous
extent, but in the limited forests of Alsace the evil was cured by the fact
that the caterpillars, having eaten up all the foliage, then died of
starvation.[300] _Teara melanosticta_ is said to produce columns of
processionary caterpillars in Australia.
FAM. 33. HYPSIDAE (or AGANAIDAE).—A family of comparatively small extent,
confined to the tropical and sub-tropical regions of the Eastern
hemisphere. The colours are frequently buff and grey, with white streaks on
the outer parts of the wings. We have nothing very like them in the
European fauna, our species of _Spilosoma_ are perhaps the nearest
approach. In _Euplocia_ the male has a pouch that can be unfolded in front
of the costa at the base of the anterior wing; it is filled with very long,
peculiar, hair-like scales growing from the costal margin; both sexes have
on each side of the second abdominal segment a small, projecting structure
that may be a sense-organ. The female is more gaily coloured than the male.
FAM. 34. ARCTIIDAE.—With the addition recently made to it of the formerly
separate family Lithosiidae, Arctiidae has become the most extensive family
of the old Bombycid series of moths, comprising something like 500 genera
and 3000 species. Hampson recognises four sub-families—Arctiinae,
Lithosiinae, {409}Nolinae, Nycteolinae,—to which may be added others from
America—Pericopinae, Dioptinae, Ctenuchinae; these sub-families being
treated as families by various authors. The sub-family Arctiinae includes
our Tiger- and Ermine-moths, and a great many exotic forms of very diverse
colours and patterns; the species of this division are, on the whole,
probably more variable in colour and markings than in any other group of
Lepidoptera. There are many cases of great difference of the sexes; in the
South American genus _Ambryllis_ the male is remarkable for its hyaline
wings with a few spots; while the female is densely scaled, and very
variegate in colour. There are some cases (the South European genus
_Ocnogyna_) where the female is wingless and moves but little, while the
male flies with great rapidity. _Epicausis smithi_, from Madagascar, one of
the most remarkable of moths, is placed in this division of Arctiidae; it
is of a tawny colour, variegate with black; the abdomen of this latter
colour is terminated by a large tuft of long scarlet hairs; the Insect has
somewhat the appearance of a Hummingbird-hawkmoth. _Ecpantheria_ is an
extensive genus of tropical American moths (having one or two species in
North America), of black and white or grey colours, with very complex
markings; the male in some species has a part of the hind wing produced as
a tail, or lobe, of a different colour.
The sub-family Pericopinae are almost peculiar to South America (two
species of _Gnophaela_ exist in North America); some of this sub-family
bear a great resemblance to Heliconiid butterflies.
The Dioptinae are likewise American moths of diurnal habits, and many of
them bear a striking resemblance to the Ithomiid butterflies they associate
with when alive.
The sub-family Lithosiinae is of great extent; our native "Footmen" give a
very good idea of it; the moths are generally of light structure, with
long, narrow front wings; a simple system of yellow and black colour is of
frequent occurrence. Many of this group feed in the larval state on
lichens. Hampson includes in this group the Nyctemeridae—light-bodied
diurnal moths, almost exclusively of black and white colours, of Geometrid
form, frequently treated as a distinct family.
The sub-family Nolinae is a small group of rather insignificant Insects, in
appearance like Pyralids or Geometrids; four or five {410}species are
native in Britain. Packard maintains the family Nolidae as distinct.[301]
The sub-family Nycteolinae consists of a few small moths the position of
which has always been uncertain; _Nycteola_ (better known as
_Sarrothripus_), _Halias_, and _Earias_ are all British genera that have
been placed amongst Tortrices, to which they bear a considerable
resemblance. _Sarrothripus_ is at present placed by Hampson in Noctuidae,
by others in Lithosiidae, by Meyrick in Arctiidae. The sub-family forms the
family Cymbidae of Kirby;[302] it includes at present only about 70
species, all belonging to the Eastern hemisphere. Two types of larvae are
known in it: one bare, living exposed on leaves; the other, _Earias_,
hairy, living among rolled-up leaves. _Halias prasinana_ is known from the
testimony of numerous auditors to produce a sound when on the wing, but the
_modus operandi_ has not been satisfactorily ascertained. Sound-production
seems to be of more frequent occurrence in Arctiidae than it is in any
other family of Lepidoptera; _Dionychopus niveus_ produces a sound by, it
is believed, friction of the wings. In the case of the genera _Setina_ and
_Chelonia_ the process is said to be peculiar to the male sex: Laboulbène
believes it to proceed from drum-like vesicles situate one on each side of
the base of the metathorax.[303]
FAM. 35. AGARISTIDAE.—An interesting assemblage of moths, many of them
diurnal and of vivid colours, others crepuscular. There is considerable
variety of appearance in the family, although it is but a small one, and
many of its members remind one of other and widely separated families of
Lepidoptera. The style and colour of the Japanese _Eusemia villicoides_ are
remarkably like our _Arctia villica_. In some forms the antennae are
somewhat thickened towards the tip and hooked, like those of the Skipper
butterflies. The family consists at present of about 250 species, but we
doubt its being a sufficiently natural one. It is very widely distributed,
with the exception that it is quite absent from Europe and the
neighbourhood of the Mediterranean Sea. In North America it is well
represented. The larvae, so far as known, are not very remarkable; they
have some lateral tufts of hair, as well as longer hairs scattered over the
body.
{411}The male of the Indian _Aegocera tripartita_ has been noticed to
produce a clicking sound when flying, and Sir G. Hampson has shown[304]
that there is a peculiar structure on the anterior wing; he considers that
this is rubbed against some spines on the front feet, and that the sound is
produced by the friction. Though this structure is wanting in the
acknowledged congeners of _A. tripartita_, yet it occurs in a very similar
form in the genus _Hecatesia_, already noticed under Castniidae.
FAM. 36. GEOMETRIDAE (_Carpets_, _Pugs_, _etc._)—This very extensive family
consists of fragile moths, only a small number being moderately stout
forms; they have a large wing-area; the antennae are frequently highly
developed in the males, but on this point there is much diversity. Either
the frenulum or the proboscis is absent in a few cases. The caterpillars
are elongate and slender, with only one pair of abdominal feet—placed on
the ninth segment—in addition to the anal pair, or claspers. They progress
by moving these two pairs of feet up to the thoracic legs, so that the body
is thrown into a large loop, and they are hence called Loopers or
Geometers. The family is universally distributed, and occurs even in remote
islands and high latitudes; in Britain we have about 270 species. The
family was formerly considered to be closely connected with Noctuidae, but
at present the opinion that it has more intimate relations with the
families we have previously considered is prevalent. Packard considers it
near to Lithosiidae, while Meyrick merely places the six families, of which
he treats it as composed, in his series Notodontina. Hampson adopts
Meyrick's six families as subfamilies, but gives them different names,
being in this respect more conservative than Meyrick, whose recent revision
of the European forms resulted in drastic changes in nomenclature.[305]
This classification is based almost exclusively on wing-nervuration. The
number of larval legs and the consequent mode of walking is one of the most
constant characters of the group; the few exceptions that have been
detected are therefore of interest. _Anisopteryx aescularia_ has a pair of
undeveloped feet on the eighth segment, and, according to Meyrick, its
allies "sometimes show rudiments of the other two pairs." The larva of
_Himera {412}pennaria_ is said to have in early life a pair of imperfect
feet on the eighth segment, which disappear as the larva approaches
maturity.
[Illustration: Fig. 202—Larva of _Amphidasis betularia_, reposing on a
rose-twig. × 1. Cambridge.]
The position of the abdominal feet and claspers throws the holding power of
the larva to the posterior part of the body, instead of to the middle, as
in other caterpillars. This, combined with the elongate form, causes these
larvae when reposing to assume attitudes more or less different from those
of other larvae; holding on by the claspers, some of these Insects allow
all the anterior parts of the body to project in a twig-like manner. The
front parts are not, however, really free in such cases, but are supported
by a thread of silk extending from the mouth to some point near-by. Another
plan adopted is to prop the front part of the body against a twig placed at
right angles to the supporting leaf, so that the caterpillar is in a
diagonal line between the two (Fig. 202). Other Geometers assume peculiar
coiled or spiral attitudes during a whole or a portion of their lives; some
doing this on a supporting object—leaf or twig—while others hang down
(_Ephyra pendularia_). Certain of the larvae of Geometridae vary in colour,
from shades of brown to green; there is much diversity in this variation.
In some species it is simple variation; in others it is dimorphism, _i.e._
the larvae are either brown or green. In other cases the larvae are at
first variable, subsequently dimorphic. In _Amphidasis betularia_ it would
appear that when the larva is hatched the dimorphism is potential, and that
the future colour, whether {413}green or brown, is settled by some
determining condition during the first period of larval life and cannot be
subsequently modified.[306] According to Poulton, the dark tint is due in
_A. betularia_ to colouring matter in the skin or immediately below it, and
the green tint to a layer of fat between the hypodermis and the superficial
muscles; this layer being always green, but more brightly green in the
larvae that are of this colour externally. Much discussion has occurred
about these larval attitudes and colours, and it seems probable that
Professor Poulton has overrated the value of protection from birds, mammals
and entomologists; the chief destroying agents being other than these, and
not liable to be thus deceived, even if the vertebrates are. In some cases
such resemblance as undoubtedly exists is not made the best use of. The
larva shown in figure 202 bore a wonderful resemblance, when examined, to
the rose-twigs it lived on, but the effect of this as a concealing agent
was entirely destroyed by the attitude; for this, being on different lines
to those of the plant, attracted the eye at once. This larva, and we may
add numerous other larvae, could have been perfectly concealed by adopting
a different attitude, but never did so; the position represented being
constantly maintained except while feeding.
In some species of this family the adult females are without wings, or have
them so small that they can be of no use for flight. This curious condition
occurs in various and widely-separated groups of the Geometridae; and it
would be naturally supposed to have a great effect on the economy of the
species exhibiting it, but this is not the case. Some of the flightless
females affect the highest trees and, it is believed, ascend to their very
summits to oviposit. It has been suggested that they are carried up by the
winged males, but this is probably only an exceptional occurrence; while,
as they are known to be capable of ascending with rapidity by means of
crawling and running, it may be taken for granted that this is the usual
method with them. Some of these wingless females have been found in numbers
on gas-lamps, and are believed to have been attracted by the light, as is
the case with very many of the winged forms.[307] {414}Neither is the
geographical distribution limited by this inferior condition of the most
important of the organs of locomotion, for _Cheimatobia brumata_ (the
Winter-moth) one of the species with flightless female, is a common and
widely distributed Insect in Europe and North America.
Although the classification of this family is based almost entirely on
wing-nervuration, yet there are some divisions of the Geometridae in which
this character is remarkably variable, certain individuals frequently
exhibiting considerable abnormality.[308] _Amphidasis betularia_ is
believed to have changed its variation considerably in the course of the
last fifty years. Previous to that time a black variety of the species was
unknown, but it has now become common; and it is believed that other
species of Geometridae are in process of exhibiting a similar
phenomenon.[309]
FAM. 37. NOCTUIDAE (_Owlet-Moths_, _Eulen_ of the Germans).—This very
extensive assemblage consists of moths rarely seen in the day-time, of
generally sombre colours, with antennae destitute of remarkable
developments in the male (except in a small number of forms); proboscis and
frenulum both present; a complex sense-organ on each side of the body at
the junction of the metathorax and abdomen. The number of species already
known can scarcely be less than 8000; owing to their large numbers and the
great general resemblance of the forms, their classification is a matter of
considerable difficulty. Although the peculiar structure at the base of the
thorax was long since pointed out, it has never received any thorough
investigation. Few other remarkable structures have yet been discovered:
the most interesting is perhaps the peculiarity in the hind wings of the
males of certain Ommatophorinae recently pointed out by Sir G. F.
Hampson[310]: in the genera _Patula_ and _Argiva_ the form of the hind
wings is normal in the females, but in the male the anterior one-half of
each of these wings is aborted, and the position of the nervures changed;
this condition is connected with the development of a glandular patch or
fold on the wing, and is remarkable as profoundly affecting a structure
which is {415}otherwise so constant that the classification of the family
is largely based on it.
[Illustration: Fig. 203—_Brephos notha._ Larva, newly hatched. Britain.]
The larvae are as a rule destitute of the remarkable adornments of hairs
and armatures of spines that are so common in many of the families we have
previously considered; they are fond of concealing themselves during the
day and coming out at night to feed; many of them pass most of their time
at, or beneath, the surface of the ground, finding nourishment in roots or
the lower parts of the stems of plants; this is notably the case in the
genus _Agrotis_, which is perhaps the most widely distributed of all the
genera of moths. Such caterpillars are known as Cut-worms in North
America.[311] The great resemblance, _inter se_, of certain of these
Cut-worms, much astonished the American naturalist Harris, who found that
larvae almost perfectly similar produced very different moths. The majority
of Noctuid larvae have the usual number of legs, viz., three pairs of
thoracic legs, four pairs of abdominal feet and the terminal claspers. In
some divisions of the family there is a departure from this arrangement,
and the abdominal feet are reduced to three, or even to two, pairs. One or
two larvae are known—e.g. _Euclidia mi_—in which the claspers have not the
usual function, but are free terminal appendages. When the abdominal legs
are reduced in number (_Plusia_, e.g.) the larvae are said to be
Half-loopers, or Semi-loopers, as they assume to some extent the peculiar
mode of progression of the Geometrid larvae, which are known as Loopers. In
the case of certain larvae, e.g. _Triphaena_, that have the normal number
of feet, it has been observed that when first hatched, the one or two
anterior pairs of the abdominal set are ill developed, and the larvae do
not use them for walking. This is the case with the young larva of our
British _Brephos notha_ (Fig. 203). Subsequently, however, this larva
undergoes a considerable change, and appears in the form shown in Fig. 204.
This interesting larva joins together two or three {416}leaves of aspen and
lives between them, an unusual habit for Noctuid larvae. When about to
pupate it bores into bark or soft wood to change to a pupa, Fig. 205; the
specimen represented closed the hole of entry by placing two separate doors
of silk across the burrow, as shown at _d_. The anal armature of this pupa
is terminated by a curious transverse process. The systematic position of
this interesting Insect is very uncertain: Meyrick and others associate it
with the Geometridae.
[Illustration: Fig. 204—_Brephos notha._ Adult larva.]
[Illustration: Fig. 205 —_Brephos notha._ A, Pupa, ventral aspect; B,
extremity of body, magnified; C, the pupa in wood; _d_, diaphragms
constructed by the larva.]
The larva of _Leucania unipunctata_ is the notorious Army-worm that commits
great ravages on grass and corn in North America. This species sometimes
increases in numbers to a considerable extent without being observed, owing
to the retiring habits of the larvae; when, however, the increase of
numbers has been so great that food becomes scarce, or for some other
cause—for the scarcity of food is supposed not to be the only reason—the
larvae become gregarious, and migrate in enormous swarms: whence its
popular name. The Cotton-worm, _Aletia xylinae_ is even more notorious on
account of its ravages. Riley states[312] that in bad years the mischief it
commits on the cotton crop causes a loss of £6,000,000, and that for a
period of fourteen successive years the annual loss averaged about
£3,000,000. This caterpillar strips the cotton plants of all but their
branches. It is assisted in its work by another highly destructive Noctuid
caterpillar, the Boll-worm, or larva of _Heliothis armigera_, which bores
into the buds and pods. This {417}latter Insect attacks a great variety of
plants, and has a very wide distribution, being found even in England,
where happily it is always a rare Insect.
In Britain, as well as in parts of Northern Europe, a Noctuid moth,
_Charaeas graminis_, occasionally increases to an enormous extent: its
larva is called the Hill-grub and lives on the grass of pastures,
frequently doing great damage in hill-lands. The increase of this moth
seems to take place after the manner of an epidemic; a considerable number
of years may pass during which it is scarcely seen, and it will then appear
in unusual numbers in widely separated localities. This moth lays a large
number of eggs, and is not completely nocturnal in habits; sometimes it may
be seen on the wing in great numbers in the hottest sunshine, and it has
been noticed that there is then a great disproportion of the sexes, the
females being ten or twenty times as numerous as the males. In Australia,
the Bugong moth, _Agrotis spina_, occurs in millions in certain localities
in Victoria: this moth hibernates as an imago, and it formerly formed, in
this instar, an important article of food with the aborigines. The powers
of increase of another Noctuid moth—_Erastria scitula_—are of great value.
Its habits have been described by Rouzaud.[313] On the shores of the
Mediterranean the larva of this little moth lives on a
Scale-Insect—_Lecanium oleae_—that infests the peach; and as the moth may
have as many as five generations in a year, it commits laudable havoc with
the pest. The larva is of remarkable form, very short and convex, with
small head, and only two pairs of abdominal feet. The scale of the
_Lecanium_ is of larger size than is usual in that group of Insects, and
the young larva of the _Erastria_ buries itself, as soon as hatched, in one
of the scales; it destroys successively numerous scales, and after having
undergone several moults, it finds itself provided, for the first time,
with a spinneret, when, with the aid of its silk, it adds to and adapts a
Coccid scale, and thus forms a portable habitation; this it holds on to by
means of the pair of anal claspers, which are of unusual form. The case is
afterwards subjected to further alteration, so that it may serve as a
protection to the creature when it has changed to a pupa. This moth is said
to be free from the attacks of parasites, and if this be the case it is
probable that its increase is regulated by the fact that {418}when the
creature becomes numerous it thus reduces the food supply, so that its own
numbers are afterwards in consequence diminished.
One of the most remarkable genera of British Noctuidae is _Acronycta_,[314]
the larvae of which exhibit so much diversity that it has been suggested
that the genus should be dismembered and its fragments treated as allied to
several different divisions of moths. There are many points of interest in
connection with the natural history of these _Acronycta_. _A. psi_ and _A.
ridens_ are practically indistinguishable as moths, though the larvae are
easily separated: the former species is said to be destroyed to an amazing
extent by parasites, yet it remains a common Insect. The genus _Apatela_ is
very closely allied to _Acronycta_, and Harris says that "_Apatela_
signifies deceptive, and this name was probably given to the genus because
the caterpillars appear in the dress of Arctians and Liparians, but produce
true owlet-moths or Noctuas."[315] The species of another British genus,
_Bryophila_, possess the exceptional habit of feeding on lichens. Some of
the American group Erebides are amongst the largest Insects, measuring
seven or eight inches across the expanded wings.
The Deltoid moths are frequently treated as a distinct family, Deltoidae,
perhaps chiefly because of their resemblance to Pyralidae. At present,
however, they are considered to be separated from Noctuidae by no valid
characters.
FAM. 38. EPICOPEIIDAE.—The genus _Epicopeia_ consists of only a few moths,
but they are amongst the most extraordinary known: at first sight they
would be declared without hesitation to be large swallow-tail butterflies,
and Hampson states that they "mimic" the Papilios of the _Polyxenus_ group.
Very little is known about these extremely rare Insects, but the larva is
stated, on the authority of Mr. Dudgeon, to surpass the moths themselves in
extravagance; to be covered with long processes of snow-white
efflorescence, like wax, exuded from the skin, and to "mimic" a colony of
the larva of a Homopterous Insect. Some ten species of this genus are known
from Java, India, China, and Japan. In this family there is said to be a
rudimentary frenulum, but it is doubtful whether the hairs that have given
rise to this definition really justify it.
{419}FAM. 39. URANIIDAE.—A family of small extent, including light-bodied
moths with ample wings and thread-like antennae; most of them resemble
Geometridae, but a few genera, _Urania_ and _Nyctalemon_, are like
Swallow-tail butterflies and have similar habits. The Madagascar moth,
_Chrysiridia madagascariensis_ (better known as _Urania rhipheus_), is a
most elegant and beautiful Insect, whose only close allies (except an East
African congener) are the tropical American species of _Urania_, which were
till recently treated as undoubtedly congeneric with the Madagascar moth.
The family consists of but six genera and some sixty species. The question
of its affinities has given rise to much discussion, but on the whole it
would appear that these Insects are least ill-placed near Noctuidae.[316]
The larva of the South American genus _Coronidia_ is in general form like a
Noctuid larva, and has the normal number of legs; it possesses a few
peculiar fleshy processes on the back. A description of the larva of
_Chrysiridia madagascariensis_ has been widely spread; but according to
Camboué,[317] the account of the metamorphoses, first given by Boisduval,
is erroneous. The larva, it appears, resembles in general form that of
_Coronidia_, and has sixteen feet; it is, however, armed with long,
spatulate black hairs; it changes to a pupa in a cocoon of open network.
[Illustration: Fig. 206—Abdomen of _Chrysiridia madagascariensis_. A,
Horizontal section showing the lower part of the male abdomen: 1, first
segment; 2, spiracle of second segment; 4-8, posterior segments. B, the
abdomen seen from the side, with the segments numbered. The section is that
of an old, dried specimen.]
In all the species of this family we have examined, we have noticed the
existence of a highly peculiar structure that seems hitherto to have
escaped observation. On each side of the second abdominal segment there is
an ear-like opening (usually {420}much concealed by overlapping scales),
giving entrance to a chamber in the body; this chamber extends to the
middle line, being separated from its fellow by only a thin partition. At
its anterior and lateral part there is a second vesicle-like chamber,
formed by a delicate membrane that extends as far forwards as the base of
the abdomen. There can be little doubt that this is part of some kind of
organ of sense, though it is much larger than is usual with Insect
sense-organs.
FAM. 40. EPIPLEMIDAE.—Under this name Hampson has assembled certain
Geometroid moths, some of them placed previously in Chalcosiidae, some in
Geometridae. They form a varied group, apparently closely allied to
Uraniidae, and having a similar peculiar sense-organ; but are distinguished
by the presence of a frenulum. The larva seems to be like that of
Uraniidae.
FAM. 41. PYRALIDAE.—This division is to be considered rather as a group of
families than as a family; it includes a very large number of small or
moderate-sized moths of fragile structure, frequently having long legs;
antennae simple, only in a few cases pectinate; distinguished from
Noctuidae and all the other extensive divisions of moths by the peculiar
course of the costal nervure of the hind wing, which either keeps, in the
middle of its course, near to the sub-costal or actually unites with it,
subsequently again separating. Members of the Pyralidae are found in all
lands; in Britain we have about 150 species. The larvae are usually nearly
bare, with only short, scattered setae, and little coloration; they have
most varied habits, are fond of concealment, and are very lively and abrupt
in movement, wriggling backwards as well as forwards, when disturbed; a
cocoon is formed for the metamorphosis.
The family as a whole consists of Insects of unattractive appearance,
although it contains some very elegant and interesting moths and numerous
forms of structural interest. In the genus _Thiridopteryx_ little
transparent spaces on the wings occur as a character peculiar to the males;
the spaces are correlative with a greater or less derangement of the
wing-nervures. In some other forms there is a remarkable retinaculum,
consisting of large scales, and this, too, is connected with a distortion
of the wing-nervures. The Pyralidae—Pyralites of Ragonot,[318] Pyralidina
of {421}Meyrick—have recently been revised by two naturalists of
distinction almost simultaneously; unfortunately their results are
discrepant, Meyrick including Pterophoridae and Orneodidae, and yet
admitting in all only eight families; while Ragonot does not include the
two groups named, but defines seventeen tribes of the two
families—Pyralidae and Crambidae—that he admits.
The Pyraustidae of Meyrick is an enormous division including the
Hydrocampidae and Scopariidae of many authors, as well as the Pyraustinae
proper and a small group of Ragonot's, the Homophysinae. The division
Scopariinae is believed to be amongst the "most ancient" of Lepidoptera;
the food of the larvae consists of moss and lichens. This group is widely
distributed, being richly represented in Australia, New Zealand, and the
Hawaiian Islands, as well as in Europe; and probably really occurs wherever
their food-plants exist accompanied by a tolerable climate. The statistics
of the distribution of this group, so far as at present known, have been
furnished by Mr. Meyrick, as follows:—European region, about 25 species;
Madeira, 3; St. Helena, 6; South Africa, 2 or 3; India, 9; Malayan region,
3 or 4; Australia, 24; New Zealand, 64; Hawaiian Islands, 50; North
America, 17 (one of them European); South America, 10. The
Hydrocampinae—the China-marks—are of great interest, as being amongst the
few forms of Lepidoptera adapted for aquatic life. It is believed that all
their larvae are aquatic, though of only a few is there much known. The
diversity amongst these forms is of considerable interest. The habits of
_Hydrocampa nymphaeata_ were long since described by Réaumur, and have more
recently been dealt with by Buckler,[319] W. Müller[320] and Prof.
Miall.[321] Although there are some discrepancies in their accounts, due we
believe to the observations being made at different periods of the life and
under somewhat different circumstances, yet the account given by Müller is
we feel no doubt substantially correct. The larvae when hatched mine in the
leaves of a water-plant for a short time—thirty hours to three days
according to Buckler—and are completely surrounded by water, which
penetrates freely into their burrows; at this period the caterpillar
breathes by its skin, the spiracles being very small, and the tubes leading
from {422}them closed and functionless. After this brief period of mining
life, the larva moults and then constructs a habitation by cutting a piece
out of a leaf, and fastening it to the under side of another leaf; it is
thus provided with a habitation, but it is one into which the water freely
enters, and the respiratory apparatus remains in the state we have
described. The Insect passes through several moults, and then hibernates in
the water. On its revival in the spring a change occurs, and the larva
constructs a portable, or we should rather say free, habitation out of two
large pieces of leaf of lens-shape, fastened together at the edges; but the
larva has some method of managing matters so that the water can be kept out
of this house; thus the creature lives in air though immersed in the water.
A correlative change occurs in the structure of the skin and tracheal
system. The former becomes studded with prominent points that help to
maintain a coat of air round the Insect, like dry velvet immersed in water;
the spiracles are larger than they were, and they and the tracheal tubes
are open. One or two moults take place and the creature then pupates. There
is a good deal of discrepancy in the accounts of this period, and it seems
probable that the pupa is sometimes aerial, sometimes aquatic. Buckler's
account of the formation of the case shows that the larva first cuts off,
by an ingenious process, one piece of leaf, leaving itself on this, as on a
raft; this it guides to a leaf suitable for a second piece, gets the raft
underneath, and fastens it with silk to the upper portion, and then severs
this, leaving the construction free; afterwards the larva goes through a
curious process of changing its position and working at the two extremities
of the case, apparently with the object of making it all right as regards
its capacity for including air and keeping out water. He believes that
Réaumur was correct in his idea that the larva regulates the admission of
air or of water to the case in conformity with its needs for respiration.
Müller calls special attention to the great changes in habit and in the
structure of the integument during the life of this larva; but the reader
will gather from what we relate as to various terrestrial Lepidopterous
larvae, that these phenomena are not very dissimilar from what frequently
take place in the latter; a change of habits at some particular moult,
accompanied by great changes in the integument, and even in the size of the
stigmata, being of frequent occurrence.
{423}The larva of _Nymphula stagnata_, a close ally of _H. nymphaeata_, has
aquatic habits of a somewhat similar but simpler nature; while _N._
(_Paraponyx_) _stratiotata_ is very different. This larva is provided with
eight rows of tufts of flexible branchiae, occupying the position of the
spots or setigerous warts usual in caterpillars, and reminding one of the
spines of certain butterfly-larvae, though they are undoubtedly respiratory
filaments. These caterpillars protect themselves by forming silken webs or
cases, or by adopting the case of some other larva, and are in the habit of
holding on by the anal claspers, and rapidly and energetically moving the
anterior parts of the body in an undulating fashion. The spiracles exist,
but are functionless. The pupa lives under water, and has no branchiae; but
three of the pairs of abdominal spiracles are open, and project from the
body. Müller informs us that in a Brazilian _Paraponyx_ these three pairs
of spiracles were already large in the larva, though the other pairs were
very small, or absent. He considers that the moth of this species descends
beneath the water of a rapid stream, and fastens its eggs on the stems of
plants therein. _Cataclysta lemnata_ lives in a case of silk with leaves of
duckweed attached to it, or in a piece of a hollow stem of some aquatic
plant; it is believed to breathe, like _H. nymphaeata_, at first by the
integument and subsequently by open stigmata; but particulars as to how it
obtains the requisite air-supply are not forthcoming: the aquatic pupa
breathes by three large abdominal spiracles like _Paraponyx_.
Musotimidae[322] is a small group of two or three genera found in Australia
and Polynesia; and the Tineodidae also consist of only two Australian
genera. Siculodidae is likewise a small Antarctic group, placed by Meyrick
in Pyralidina; but his view is not accepted by Snellen and Ragonot.
Epipaschiinae (formerly treated as a separate family) and Endotrichiinae
are, according to Meyrick, subdivisions of the family Pyralidae proper, an
enormous group of more than 100 genera. The Chrysauginae consist chiefly of
American forms, and have not been treated by Meyrick; some of this group
have been classed with Tortricidae or Deltoidae on account of the
undulating costa of the front wings and the long, peculiar palpi. The
Galleriidae are a small group including Insects that live in bees'-nests,
and feed on the wax {424}etc.; others eat seeds, or dried vegetable
substances. Three out of our five British species of this family occur
(usually gregariously) in bee-hives, and have the peculiar habit of
spinning their cocoons together. The mass of common cocoons formed in this
manner by _Aphomia sociella_ is remarkably tough and enduring; portions of
it are not infrequently picked up, and as the cocoons are of a peculiar
tubular form their nature gives rise to some perplexity.
Phycitidae[323] is another very large assemblage of Insects with very
diverse habits. The frenulum and retinaculum are similarly formed in the
two sexes: the males frequently have the basal-joint of the antennae
swollen; hence the term "Knot-horns" applied by collectors to these moths.
The larvae of the species of _Ephestia_ infest groceries, and most children
have become to a slight extent acquainted with them amongst dried figs;
that of _E. kuehniella_ has become very injurious in flour-mills, its
enormous increase being due in all probability to the fact that the
favourable and equable temperature maintained in the mills promotes a rapid
succession of generations, so that the Insect may increase to such an
extent as to entirely block the machinery. Many of the Phycitidae feed on
the bark of trees in galleries or tunnels constructed partially of silk. A
very peculiar modification of this habit in _Cecidipta excoecaria_ has been
described by Berg.[324] In Argentina this Insect takes possession of the
galls formed by a _Chermes_ on _Excoecaria biglandulosa_, a Euphorbiaceous
tree. The female moth lays an egg on a gall, and the resulting larva bores
into the gall and nourishes itself on the interior till all is eaten except
a thin external coat; the caterpillar then pupates in this chamber. The
galls vary in size and shape, and the larva displays much constructive
ability in adapting its home to its needs by the addition of tubes of silk
or by other modes. Sometimes the amount of food furnished by the interior
of the gall is not sufficient; the larva, in such cases, resorts to the
leaves of the plant for a supplement, but does not eat them in the usual
manner of a caterpillar; it cuts off and carries a leaf to the entrance of
its abode, fastens the leaf there with silk, and then itself entering,
feeds, from the interior, on the food it has thus acquired. Another
Phycitid, _Dakruma coccidivora_ is very {425}beneficial in North America by
eating large Scale-Insects of the _Lecanium_ group, somewhat after the
fashion of _Erastria scitula_; it does not construct a case, but shelters
itself when walking from one scale to another by means of silken tubes; it
suffers from the attacks of parasites.[325] Oxychirotinae, an Australian
group, is interesting because, according to Meyrick, it possesses forms
connecting the Pterophoridae with the more normal Pyralids.
Crambidae, or Grass-moths, are amongst the most abundant Lepidoptera in
this country, as they include the little pale moths that fly for short
distances amongst the grass of lawns and pastures; they fold their wings
tightly to their body, and have a head pointed in front, in consequence of
the form and direction of the palpi. They sit in an upright position on the
stems of grass, and it has been said that this is done because then they
are not conspicuous. Perhaps: but it would be a somewhat difficult
acrobatic performance to sit with six legs across a stem of grass. The
larvae are feeders on grass, and construct silken tunnels about the roots
at or near the surface. The Ancylolominae are included in Crambidae by
Meyrick and Hampson. Schoenobiinae[326] are included by Meyrick in
Pyraustidae, but this view appears not to meet with acceptance, and the
group is more usually associated with the Crambidae. Most writers place the
anomalous genus _Acentropus_ as a separate tribe, but it is associated by
both Meyrick and Hampson with _Schoenobius_. This Insect is apparently the
most completely aquatic of all the Lepidoptera, and was for long associated
with the Trichoptera in consequence of its habits and of the scaling of the
wings being of a very inferior kind. The males may sometimes be found in
large numbers fluttering over the surface of shallow, but large, bodies of
water; the females are rarely seen, and in some cases have no wings, or
have these organs so small as to be useless. The female, it would appear,
comes quite to the surface for coupling, and then takes the male beneath
the water. The larvae have the usual number of Lepidopterous feet, and
apparently feed on the leaves of plants below water just as Lepidopterous
larvae ordinarily do in the air.[327] They have no trace of gills, and
their {426}mode of respiration is unknown. A great deal has been written
about these Insects, but really very little is known. They are abundant,
though local in many parts of North and Central Europe; some of the females
have, as we have said, abbreviated wings, but how many species there are,
and whether the modifications existing in the development of the wings are
constant in one species or locality, are unknown as yet.
FAM. 42. PTEROPHORIDAE[328] (_Plume-moths_).—Elegant Insects of small size,
usually with the wings divided (after the fashion of a hand into fingers)
so as to form feathers: the extent of this division is diverse, but the
hind wings are more completely divided than the front, which indeed are
sometimes almost entire. The group is placed by Meyrick in his Pyralidina,
but there are many entomologists who look on it as distinct. It consists of
two sub-families, Agdistinae and Pterophorinae, that have been treated as
families by many entomologists. The Agdistinae (of which we have a British
representative of the only genus _Agdistes_) have the wings undivided.
Pterophorinae have the hind wings trifid or (rarely) quadrifid, the front
wings bifid or (rarely) trifid. The larvae of the Pterophorinae are
different from those of Pyralidae, being slow in movement and of heavy
form, covered with hair and living exposed on leaves; the pupae are highly
remarkable, being soft, coloured somewhat like the larvae, and also hairy
like the larvae, and are attached somewhat after the manner of
butterfly-pupae by the cremaster: but in some cases there is a slight
cocoon. There is, however, much variety in the larval and pupal habits of
the Pterophoridae, many having habits of concealment of divers kinds. We
have thirty species of these lovely Plume-moths in Britain. The family is
widely distributed, and will probably prove numerous in species when the
small and delicate Insects existing in the tropics are more appreciated by
collectors.
FAM. 43. ALUCITIDAE (ORNEODIDAE of Meyrick and others).—The genus _Alucita_
includes the only moths that have the front and hind wings divided each
into six feathers. Species of it, though not numerous, occur in various
regions. The larva and pupa are less anomalous than those of the
Pterophoridae, though the imago is more anomalous. The caterpillar of our
British _A. polydactyla_ feeds on the flower-buds of honey-suckle, and
forms a {427}cocoon. The moth with wings expanded is about an inch across,
and is a lovely object. It is not rare, though seldom numerous.
FAM. 44. TORTRICIDAE.—Moths of small size, with a rather ample wing area,
with the wing-fringes never as long as the wings are wide (long across),
the hind wings without a pattern: the anterior nervure on the hind wings is
simply divergent from that next to it, and the internal nervure, 1b, is
very evidently forked at the base. The larvae inhabit their food, which may
be rolled up or twisted leaves, or the interior of fruits and herbs, or
galls, or even roots; they exhibit less diversity than is usual in other
large series of moths; all have the normal complement of sixteen legs. This
group is a very extensive one, but is much neglected owing to the great
difficulties attending its study; it is not recognised in Hampson's Table
of families given on p. 370, being there merged in Tineidae. It appears,
however, to be a really natural group, and it is not desirable to merge it
in the sufficiently enormous assemblage of the Tineidae till this has been
shown to be necessary by the light of a greater knowledge of the external
anatomy than we possess at present. The term Microlepidoptera is frequently
met with in entomological literature, and should, we think, be confined to
the two series Tortricidae and Tineidae. The Pterophoridae, and even the
Pyralidae, have been, and still sometimes are, included under this term,
but at present it seems best to limit its application as is here suggested.
Three great divisions are at present recognised; these were formerly called
by Meyrick,[329] Tortricidae, Grapholithidae, Conchylidae;
subsequently,[330] he has adopted the names Tortricidae, Epiblemidae,
Phaloniadae. Lord Walsingham, who has devoted a great deal of time and
study to the elucidation of this most difficult group, has suggested[331]
that another change is desirable, and if so the nomenclature will be:—1.
Tortricidae [or Tortricinae, according to the view that may be taken as to
the group being family or sub-family]; 2. Phaloniidae [= the formerly used
name, Conchylidae]; 3. Olethreutidae [= the formerly used name
Grapholithinae = Epiblemidae, Meyr.]. We have upwards of 300 species in
Britain, nearly 200 of which belong to the last division. The name
Tortricidae refers to the habit the {428}larvae of these moths possess of
rolling up leaves, or twisting and distorting shoots and buds.
The mode in which leaves and shoots are twisted and rolled by the very
small larvae has been much discussed and is probably the result of two or
three distinct causes:—1, the immediate operations of the larva; 2, the
contraction of silk when drying; 3, changes in the mode of growth of the
parts of the vegetable, resulting from the interference of the caterpillar.
The larvae of this family that live in fruits are only too widely (we will
not say well) known. Stainton gives as the habitat of _Epinotia funebrana_,
"larva frequent in plum-pies"; the caterpillar of _Carpocapsa pomonella_
(the Codling-moth) mines in apples and pears, and its ravages are known
only too well in widely distant parts of the world where fruit-trees of
this kind are cultivated. _C. splendana_ lives in acorns and walnuts; _C.
juliana_ in Spanish chestnuts. Two, if not more, larvae live in the seeds
of Euphorbiaceous plants, and have become notorious under the name of
jumping-beans, on account of the movements they cause. As these latter show
no trace externally of being inhabited, the movements are supposed to be a
mysterious property of the seed; they are really due to its containing a
large cavity, extending, in one direction of the seed, nearly or quite from
skin to skin; in this the larva makes a movement sufficient to alter the
point of equilibrium of the quiescent seed, or as a free body to strike
some part of it. The exact nature of the movements of the larva have not,
we believe, been ascertained. There are, at least, two species of these
Insects, and two plants harbouring them, known in the United States and
Mexico, viz. _Carpocapsa saltitans_ living in the seeds of _Croton
colliguaja_ and _Grapholitha sebastianiae_ living in the seeds of
_Sebastiania bicapsularis_.
FAM. 45. TINEIDAE.—Small moths with the labial palpi more flexible and
mobile than in other moths; usually separated and pointed. Hind wings
frequently with very long fringes, the wing itself being proportionally
reduced in size, and in consequence pointed at the tip. Larvae very
diverse, almost always with habits of concealment. The series of forms
included under this head is very numerous, the British species alone
mounting up to 700, while the total described cannot be less than 4000.
This number, however, must be but a fragment of what exists, if Mr. Meyrick
be correct in supposing that a single one of the divisions of the
family—Oecophoridae—comprises 2000 species in Australia and New Zealand
alone.
{429}[Illustration: Fig. 207—_Diplosara lignivora_ (Gelechiides). Hawaiian
Islands.]
As the study of these Insects is attended with great difficulty on account
of their fragility and the minute size of the great majority, it is not a
matter for surprise that their classification is in a comparatively
rudimentary state. We shall not, therefore, deal with it here. Neither can
we attempt to give any idea of the extreme diversity in the colours, forms,
and attitudes of these small Insects. The one shown in Fig. 207, is
remarkable on account of the great accumulation of scales on the wings and
legs. As regards the pointed wings and the long fringes, we may remark that
it is probable that in many of these small forms the wings are passive
agents in locomotion; a similar condition of the wings is found in other
very minute Insects, _e.g._ Thysanoptera and Trichopterygidae; in all these
cases the framework of the wings is nearly absent: in some forms of the
Tineidae, _Opostega_, e.g. the nervules are reduced to three or four in
each wing. The variety in habits is as great as that of the external form,
and the larvae exceed in diversity those of any other group of Lepidoptera.
No doubt a corresponding amount of diversity will be discovered in the
details of structure of the perfect Insects, the anatomy of but few having
been at present investigated. _Tinea pellionella_ has two very important
peculiarities in its internal anatomy: the testes consist of four round
follicles on each side, and, contrary to the condition generally prevalent
in Lepidoptera, are not brought together in a common capsule: the two
groups are, however, not quite free (as they are in _Hepialus_), but are
connected by a loose tracheal network. Even more remarkable is the fact
also pointed out by Cholodkovsky[332] that the adult Insect possesses only
two Malpighian tubes instead of six, the normal number in Lepidoptera; in
the larva there are, however, six elongate tubes. The group of forms to
which {430}_Tinea_ belongs is remarkable for the diversity and exceptional
character of the food-habits of the larvae; species subsist on dried
camel's dung, various kinds of clothes, furs, and hair, and even about
horns of deer and horses' hoofs: one species has been found in abundance in
the hair of a live sloth, _Bradypus cuculliger_, under circumstances that
render it possible that the larva feeds on the creature's hair, though it
may feed on minute vegetable matter found in the hair. The larva of _Tinea
vastella_ is occasionally found feeding on the horns of living antelopes.
Several species of Tineidae are known to devour Scale-Insects.
_Lita solanella_ is notorious for the ravages it commits on stored
potatoes. Quite a number of species live on cryptogamic matter, or in old
wood; _Oinophila v-flavum_ feeds on the mould on the walls of cellars, and
is reputed to be injurious by occasionally also attacking the corks of
bottles containing wine. _Oecocecis guyonella_ is said to be the cause of
galls on _Limoniastrum guyonianum_, a plant that, growing in the deserts to
the south of Algeria, is a favourite food of camels, and is frequently
entirely covered with sand. The deposition of an egg by this moth is
believed by Guénée[333] to give rise to a gall in which the larva is
entirely enclosed (like the larvae of the gall-flies). Of Clothes-moths
there are at least three species widely distributed. _Trichophaga
tapetzella_ is perhaps entitled to be considered the Clothes-moth; its
caterpillar not only feeds on clothes, but spins webs and galleries amongst
them. _Tinea pellionella_ is also very common; its larva lives in a
portable case, while that of the third species, _Tineola biselliella_,
forms neither a case nor definite galleries. We have found this the most
destructive of the three at Cambridge. Clothes or valuable furs may be
completely protected by wrapping them in good sound paper in such a way
that no crevices are left at the places where the edges of the paper meet.
Garments that have become infested may be entirely cleared by free exposure
to air and sunshine.
Two species of _Tinea_ have been recorded as viviparous, viz. _Tinea
vivipara_ in Australia, and an undetermined species in South America. The
species of the genus _Solenobia_—in which the female is apterous—are
frequently parthenogenetic. The group Taleporiidae, to which this genus
belongs, is by some {431}classified with Psychidae, in which family, as we
have pointed out, one or two parthenogenetic forms are also known.
The larvae of Tineidae, though they do not exhibit the remarkable armature
found in so many of the larger caterpillars, are exceedingly diverse.[334]
Some are entirely destitute of feet (_Phyllocnistis_). Others are destitute
of the thoracic legs; _Nepticula_ is in this case, but it is provided with
an increased number of abdominal feet, in the form of more or less
imperfect ventral processes. Some mine in leaves, others live in portable
cases of various forms. Some are leaf-miners during their early life, and
subsequently change their habits by constructing a portable case. The genus
_Coleophora_ affords numerous instances of this mode of life; the habits of
these case-bearers exhibit considerable variety, and there are many points
of interest in their life-histories. Change of habit during the larval life
has already been alluded to as occurring in many Lepidoptera and is nowhere
more strikingly exemplified than in certain Tineidae. Meyrick mentions the
following case as occurring in an Australian Insect, _Nematobola
orthotricha_;[335] the larva, until two-thirds grown, is without feet, and
is almost colourless, and mines in the leaves of _Persoonia lanceolata_;
but when two-thirds grown it acquires sixteen feet, changes colour,
becoming very variegate, and feeds externally, unprotected, on the leaves.
The cases of the case-bearing Tineids are usually of small size, and do not
attract attention like those of Psychidae. A very remarkable one was
discovered by Mr. E. E. Green in Ceylon, and was at first believed to be
formed by a Caddis-worm. It has now been ascertained that the Insect
forming it is the caterpillar of _Pseudodoxia limulus_, a Tineid moth of
the group Depressariidae;[336] the case is composed of minute fragments of
moss, sand, and lichens; the anterior end is dilated into a shield-like
hood that covers and protects the anterior parts of the larva when feeding;
the food is mosses and lichens on rocks and trees. Before pupating, the
larva folds down the edges of the hood over the mouth of the tube, like an
envelope, fastening them with silk. The case is fixed to the rock or other
support and hangs there until the moth appears.
{432}[Illustration: Fig. 208—_Pronuba synthetica_. North America. A, Larva;
B, C, pupa, ventral and lateral aspects; D, female moth; E, head and part
of thorax of the female moth: _a_, labial palp; _b_, maxillary tentacle;
_c_, maxillary palp; _d_, proboscis; _e_, base of front leg. (After
Riley.)]
The family Prodoxidae consists of some Tineids, the larvae of which feed in
the pods and stems of the Yuccas of south-western North America; they have
the mouth of very unusual form (Fig. 208, E), and some of them, by aid of
this peculiar mouth, exhibit a remarkable modification of instinct. The
facts are chiefly known from the observations of Riley[337] on _Pronuba
yuccasella_, a moth living on _Yucca filamentosa_; this plant has been
introduced into our gardens in this country, where it never, we believe,
produces seed. The Yuccas are not fitted for self-fertilisation or for
fertilisation by Insect agency of an ordinary kind. The progeny of the moth
develops in the pods of the plant, and as these cannot grow until the
flowers have been fertilised, the moth has the habit of fertilising the
flowers at the time she lays her egg in the part that is to develop into
the pod, and to be the food for her own progeny. The female moth first
visits the stamens, and collects, by the aid of the {433}maxillae (which in
this sex are very remarkably formed),[338] a considerable mass of pollen,
which she holds by means of the peculiar maxillary tentacles; she then lays
an egg in the pistil, usually of some flower other than that from which she
has gathered the pollen; and after she has accomplished this act she
carefully applies the pollen she had previously collected to the pistil, so
as to secure the fertilisation of the flower and the development of the
pod.
The species of _Prodoxus_ stand in a very peculiar relation to _Pronuba_.
They also live in Yuccas, and have habits similar to those of _Pronuba_,
with the important exception that, being destitute of the requisite
apparatus, they do not fertilise the Yucca-flowers, and are thus dependent
on _Pronuba_ for the steps being taken that are necessary for the rearing
of the progeny of the two kinds of moth. Hence the name of Yucca-moth has
been bestowed on _Pronuba_, and that of "bogus Yucca-moth" on the
_Prodoxus_. The _Pronuba_ we figure is the largest and most remarkable
species of the genus and fertilises _Yucca brevifolia_; the larva is
destitute of abdominal feet, and in the pupa the spines on the back that
exist in nearly all pupae that live in stems are developed to an
extraordinary extent. The Yuccas do not flower every year, and the
Prodoxidae have a corresponding uncertainty as to their periods of
appearance, passing sometimes a year or two longer than usual in the pupal
stage.
FAM. 46. ERIOCEPHALIDAE.—This family has recently been proposed for some of
the moths formerly included in the genus _Micropteryx_.[339] They are
small, brilliant, metallic Insects, of diurnal habits, but are very rarely
seen on the wing, and it is doubtful whether they can fly much. These
little Insects are of peculiar interest, inasmuch as they differ from the
great majority of the Lepidoptera in at least two very important points,
viz. the structure of the wings and of the mouth-parts. The mouth shows
that we may consider that the Lepidoptera belong to the mandibulate
Insects, although in the great majority of them the mandibles in the final
instar are insignificant, functionless structures, or are entirely absent,
and although the maxillae are so highly adapted for the tasting of sweets
that it is difficult to recognise in them the parts usually found in the
maxilla of mandibulate Insects.
{434}[Illustration: Fig. 209.—Larva of _Eriocephala calthella_. (After
Chapman.) A, Young larva from side, × 50; B, portion of skin with a bulla
or ball-like appendage: C, abdominal foot of larva.]
_Eriocephala_ in both these respects connects the Lepidoptera with
Mandibulata: the mandibles have been shown by Walter[340] to be fairly well
developed; and the maxillae are not developed into a proboscis, but have
each two separate, differentiated—not elongated—lobes, and an elongate,
five-jointed, very flexible palpus. The moths feed on pollen, and use their
maxillae for the purpose, somewhat in the style we have mentioned in
Prodoxidae. The wings have no frenulum, neither have they any shoulder, and
they probably function as separate organs instead of as a united pair on
each side: the modification of the anterior parts of the hind wing—whereby
this wing is reduced as a flying agent to the condition of a subordinate to
the front wing—does not here exist: the hind wing differs little from the
front wing in consequence of the parts in front of the cell being well
developed. There is a small jugum. These characters have led Packard to
suggest that the Eriocephalidae should be separated from all other
Lepidoptera to form a distinct sub-Order, Lepidoptera Laciniata.[341] The
wing-characters of _Eriocephala_ are repeated—as to their main features—in
Hepialidae and Micropterygidae; but both these groups differ from
_Eriocephala_ as to the structure of the mouth-parts, and in their
metamorphoses. Although _Eriocephala calthella_ is one of our most abundant
moths, occurring in the spring nearly everywhere, and being easily found on
account of its habit of sitting in buttercup-flowers, yet its metamorphoses
were till recently completely unknown. Dr. Chapman has, however, been able
to give us some information as to the habits and structure of the larvae,
in both of which points the creature is most interesting. The eggs and
young larvae are "quite {435}unlike our ideas of a Lepidopterous Insect;"
the former have a snowy or mealy appearance, owing to a close coating of
minute rods standing vertically on the surface of the egg, and often tipped
with a small bulb. The larva lives amongst wet moss and feeds on the
growing parts thereof; it is not very similar to any other Lepidopterous
larva: Dr. Chapman suggests a similarity to the Slug-worms (Limacodids),
but Dyar is probably correct in thinking the resemblances between the two
are unimportant: the larva of _Eriocephala_ possesses three pairs of
thoracic legs, and eight pairs of abdominal appendages, placed on the
segments immediately following the thorax; on the under-surface of the
ninth and tenth abdominal segments there is a sucker, trifoliate in form;
this is probably really situate entirely on the tenth segment: the body
bears rows of ball-appendages, and the integument is beautifully
sculptured. The head is retractile and the antennae are longer than is
usual in caterpillars. This larva is profoundly different from other
Lepidopterous larvae inasmuch as the abdominal feet, or appendages, are
placed on different segments to what is customary, and are of a different
form. Unfortunately the pupa has not been procured, but there is some
reason for supposing that it will prove to be more like that of Tineidae
than like that of Micropterygidae.
The New Zealand genus _Palaeomicra_ is only imperfectly known. Meyrick
considers it the "most ancient" Lepidopteron yet discovered; and it would
appear that its relations are with _Eriocephala_ rather than with
_Micropteryx_. From information he has kindly given to us, we are able to
say that this moth possesses mandibles but no proboscis.
[Illustration: Fig. 210—Larva of _Micropteryx_ sp. A, Ventral view of the
larva, magnified; B, the same, with setae unduly magnified. Britain.]
FAM. 47. MICROPTERYGIDAE.—Small moths of metallic colours, without
mandibles, with elongate maxillary palpi: without frenulum: both wings with
a complex system of wing-veins: on the hind wings the area anterior to the
cell is large, and traversed by three or four elongate, parallel veins.
{436}[Illustration: Fig. 211—Pupa of _Micropteryx_ (_semipurpurella_?). A,
Dorsal aspect; B, C, D, views of head dissected off; B, profile; C,
posterior, D, anterior aspects; _m_, mandibles. Britain.]
There are no mandibles, but there is a short, imperfect proboscis. Larva
(Fig. 210) without any legs, mining in leaves. The pupa (Fig. 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 {437}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).
{438}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, {439}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.
[Illustration: 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 {440}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.
{441}[Illustration: 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 {442}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, {443}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.
[Illustration: 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 {444}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 {445}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 {446}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.
{447}[Illustration: 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 {448}"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.
[Illustration: 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 {449}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]
[Illustration: 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 {450}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 {451}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.
[Illustration: 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.