magnified; B, part of the middle of the body; _a_, prolongation of
pronotum; _b_, tegmen; _c_, wing.]
The tribe Tettigides is a very extensive group of small Acridiidae, in
which the pronotum extends backwards as a hood and covers the body, the
tegmina and wings being more or less modified. In our British species (Fig.
179) this condition does not greatly modify the appearance of the Insect,
but in many exotic species (Fig. 180) the hood assumes {301}remarkable
developments, so that the Insects have no longer the appearance of
Orthoptera. It would be impossible, without the aid of many figures, to
give an idea of the variety of forms assumed by this prothoracic expansion.
It is a repetition of what occurs in the Order Hemiptera, where the
prothoracic hoods of the Membracides exhibit a similar, though even more
extraordinary, series of monstrous forms. So great is the general
similarity of the two groups that when the genus _Xerophyllum_ (Fig. 180,
A) was for the first time described, it was treated by the describer as
being a bug instead of a grasshopper. This genus includes several species
from Africa. The curious _Cladonotus_ (Fig. 180, B) is a native of Ceylon,
where it is said to live in sandy meadows, after the fashion of our
indigenous species of _Tettix_ (Fig. 179). Very little is known as to the
habits of these curious Tettigides, but it has been ascertained that some
of the genus _Scelimena_ are amphibious, and do not hesitate to enter the
water and swim about there; indeed it is said that they prefer plants
growing under water as food. This habit has been observed both in Ceylon
and the Himalayas. The species are said to have the hind legs provided with
dilated foliaceous appendages useful for swimming.
[Illustration: FIG. 180.—Tettigides: A, _Xerophyllum simile_ Fairm.; B,
_Cladonotus humbertianus_. (After Bolivar.)]
[Illustration: FIG. 181.—A, _Mastax_ (_Erianthus_) _guttatus_, male.
Sumatra. (After Westwood.) B, profile; C, front of head.]
The tribe Mastacides includes thirty or forty species of Acridiidae with
short antennae and vertical head (Fig. 181, _Mastax guttatus_); they are
apparently all rare and little {302}known, but are widely distributed in
the tropics of the Old and New Worlds. Nothing whatever seems to be known
of their habits or of their development.
The tribe Pneumorides includes a still smaller number of species of very
aberrant and remarkable grasshoppers, of large size, with short antennae,
and with the pronotum prolonged and hood-like; they are peculiar to South
Africa. Although amongst the most remarkable of Insects, we are not able to
give any information as to their habits. It would appear from the form of
their legs that they have but little power of hopping. The species of which
we figure the female (Fig. 182) is very remarkable from the difference in
colour of the sexes. The female is so extravagantly coloured that she has
been said to look as if "got up" for a fancy-dress ball. She is of a gay
green, with pearly white marks, each of which is surrounded by an edging of
magenta; the white marks are very numerous, especially on the parts of the
body not shown in our figure; the face has magenta patches and a large
number of tiny pearly-white tubercles, each of which, when placed on a
green part, is surrounded by a little ring of mauve colour. Though the
female is certainly one of the most remarkably coloured of Insects, her
consort is of a modest, almost unadorned green colour, and is considerably
different in form. He is, however, provided with a musical apparatus, which
it is possible may be a means of pleasing his gorgeous but dumb spouse. It
consists of a series of ridges placed on each side of the inflated abdomen,
which, as we have previously (p. 200) remarked, has every appearance of
being inflated with the result of improving its resonance.
[Illustration: FIG. 182.—_Pneumora scutellaris_, female. South Africa.]
{303}The Pyrgomorphides[235] is a small tribe of about 120 described
species, two of which are found in the south of Europe (Fig. 183,
_Pyrgomorpha grylloides_). The tribe includes a number of large and curious
Insects, among them the species of _Phymateus_ and _Petasia_, with peculiar
excrescences on the pronotum and vivid colours on some parts of the body or
its appendages, which are apparently common Insects in South Africa.
The tribe Tryxalides includes a great many species of grasshoppers. In them
the front of the head joins the upper part at an acute angle (Figs. 165 and
173). This tribe and the Acridiides are the most numerous in species of the
family. To the latter belong most of the migratory locusts of the New World
(Fig. 175, _Caloptenus spretus_). A Spanish species of this tribe,
_Euprepocnemis plorans_, though provided with well-developed wings,
possesses the remarkable habit of seeking shelter by jumping into the water
and attaching itself below the surface to the stems of plants.
[Illustration: FIG. 183.—_Pyrgomorpha grylloides._ South Europe. (After
Fischer.)]
[Illustration: FIG. 184.—_Xiphocera_ (_Hoplolopha_) _asina_. S. Africa.
(After de Saussure.)]
The tribe Pamphagides[236] includes some 200 species, found chiefly in
Africa and the arid regions near the Mediterranean Sea. They are mostly
apterous forms, and this circumstance has, according to de Saussure,
exercised a marked influence on the geographical distribution of the
species. Although the tribe consists chiefly of apterous forms, several
species possess {304}well-developed wings; sometimes this is the case of
the male but not of the female. Some of the species are highly modified for
a desert life, and exhibit a great variation in the colour of the
individuals in conformity with the tint of the soil they inhabit.
_Xiphocera asina_ (Fig. 184) is thought by Péringuey to be the prey of the
extraordinary South African tiger-beetles of the genus _Manticora_.
We have already mentioned the tribe Oedipodides[237] as including most of
the species of migratory locusts of the Old World. Some striking cases of
variation in colour occur amongst the winged Oedipodides. In certain
species the hind wings may be either blue or rosaceous in colour; it is
thought that the latter is the tint natural in the species, and that it is
due to the mixture of a red pigment with the pale blue colour of the wing;
hence the blue-coloured wings are analogous to cases of albinism. But the
most remarkable fact is that this colour difference is correlative with
locality. Brunner von Wattenwyl says[238] that the blue variety of _Oe.
variabilis_ occurs only in a few localities in Europe—he mentions Vienna
and Sarepta,—and that where it occurs not a single red example can be met
with. Similar phenomena occur in other species in both Europe and North
America, and L. Bruner has suggested[239] that the phenomena in the latter
country are correlative with climatic conditions.
The group Eremobiens, a subdivision of Oedipodides, includes some of the
most interesting forms of Acridiidae. Its members have several modes of
stridulation. _Cuculligera flexuosa_ and other of the winged forms,
according to Pantel,[240] produce sounds by the friction of the middle
tibia against the wing, both of these parts being specially modified for
the purpose in the male sex. The most peculiar members of the Eremobiens
are some very large Insects, modified to an extraordinary extent for a
sedentary life in deserts and arid places. Trimen says[241] that a South
African species, _Trachypetra bufo_, which lives amongst stones, is so
coloured that he had much difficulty in detecting it, and that he noticed
in certain spots, often only a few square yards in extent, where the stones
lying on the ground were darker, lighter, or more mottled than usual, that
the individuals of the grasshopper were of a similar colour to the stones.
{305}[Illustration: FIG. 185.—_Methone anderssoni_, female. S. Africa. _a_,
Front of head; _b_, posterior leg; _c_, _d_, front and hind feet. (_c_ and
_d_ magnified, the others natural size.)]
The Insect referred to by Trimen is, we believe, the _Batrachotettix whiti_
of de Saussure. In this species the alar organs are completely absent, and
the pronotum forms a sort of hood that protects the base of the hind body.
Some of the desert Eremobiens vary so much that the differences found among
individuals of the same species {306}are said by Brunner and de Saussure to
be so great as to affect even the generic characters, and give rise to the
idea of an "uncompleted species-formation."
[Illustration: FIG. 186.—Portions of middle of the body and hind leg of
_Methone anderssoni_ ♂: _a_, femur; _b_, an inferior fold; _c_,
rattling-plate; _d_, striated surface; _e_, the adjoining sculpture; _f_,
grooved portion of tegmen. The part _e_ is really, like _d_, a portion of
the second abdominal segment, not of the third, as might be supposed from
the figure.]
_Methone anderssoni_, an inhabitant of the Karoo Desert of South Africa, is
one of the largest of the Acridiidae. A female of this species is
represented of the natural size in Fig. 185. This Insect is remarkable on
account of the complex organs for producing sound, and for the great
modification of the posterior legs (Fig. 185, _b_), which do not possess
locomotive functions, but serve as a portion of the sound-producing
apparatus, and as organs for protecting the sides of the body. This Insect
is said to be very efficient in making a noise. The sexes differ
considerably in their sound-producing organs, a portion of which are
present in the female as well as in the male (Fig. 186). Connected with the
first abdominal segment, but extending backwards on the second, there is a
peculiar swelling bearing two or three strongly raised chitinous folds
(Fig. 186, _c_). When the leg is rotated these folds are struck by some
peg-like projections situate on the inner face of the base of the femur,
and a considerable noise is thus produced. The pegs cannot be seen in our
figure. This apparatus is equally well developed in female and male. On the
second abdominal segment, immediately behind the creaking folds we have
described, there is a prominent area, densely and finely striated (Fig.
186, _d_): this is rubbed by some fine asperities on the inner part of the
femur near its base. Sound is produced by this friction on the striated
surface, the sculpture of which is abruptly contrasted with that of the
contiguous parts: these structures seem to be somewhat better developed in
the male than they are in the female, and to be phonetic, at any rate in
the former sex. {307}The male has the rudimentary tegmina (Fig. 186, _f_)
much longer than they are in the female (Fig. 185), and their prolonged
part is deeply grooved, so as to give rise to strong ridges, over which
plays the edge of the denticulate and serrate femur. There is nothing to
correspond to this in the female, and friction over the surface of this
part of the male produces a different and louder sound. There can be little
doubt that this is a phonetic structure peculiar to the male. It
approximates in situation to the sound-producing apparatus of the males of
the Stenobothri and other Acridiidae. _Methone anderssoni_ has large
tympanal organs: the small tegmina cover them up completely. In the female
the tips of the tegmina seem to be adapted for forming covering-flaps for
the tympana. In both sexes there is a sac (Fig. 186, _b_) adjoining the
structures we have mentioned, but which is not directly phonetic, though it
may be an adjunct of the apparatus.
There is no other Orthopteron in which the phonetic organs are so complex
as they are in the male of _Methone anderssoni_, and it would appear
probable that this Insect possesses the power of producing two, if not
more, distinct sounds, one in common with the female, and peculiar to this
and one or two other species; the other somewhat similar to that of other
Acridiids, and more specially developed in the male, if not absolutely
confined to it.
This Insect is of a very sedentary disposition, and when disturbed
apparently seeks safety rather by the noise it can make than by flight. Its
powers of locomotion indeed are very feeble. The alar organs are quite
rudimentary, and of no assistance whatever for movement. The hind legs seem
to be almost equally useless for this purpose; they are broader than they
are in other Acridiidae, and have different functions. When _Methone_ moves
it does so by means of the anterior four legs, on which it walks propped up
as if on stilts. When at rest the hind legs are pressed close to the body,
and the tibiae are inflexed and not seen, the creature in this position
greatly resembling a clod of earth. We know nothing of the life history of
this Insect, except that the young resemble the adult in appearance, and
are provided with the sound-producing apparatus, or some portion thereof.
The geographical distribution of the Eremobiens corresponds with that of
the Pamphagides, with two important differences, viz. that in the Old World
the former group occupies a somewhat {308}more restricted area, and that it
is represented in the New World by two peculiar North American genera,
_Haldmanella_ and _Brachystola_. _B. magna_ is an Insect nearly equal in
size to _Methone anderssoni_. Its peculiar form and movements have procured
for it in Texas and Colorado the popular names of "buffalo hopper" and
"lubber grasshopper." This Insect has not—like _Methone_—the colours of the
desert sands; it is of a green tint, with comparatively smooth body, and
during the day rests concealed under tufts of grass. It has apparently no
sound organs, though de Saussure thinks there are structures present that
are vestiges or rudiments thereof.
The family Acridiidae includes a large part of the species that make up our
meagre list of British Orthoptera. Indeed, the only native Orthoptera at
the present time sufficiently common to attract general attention are, in
addition to the earwig, the species of the genera _Stenobothrus_ and
_Gomphocerus_, whose musical instruments we have described previously. We
have eight species of these Insects. They are the little grasshoppers, so
common in our fields and gardens, the hunting of which is a source of much
amusement to children. The Insect goes off with a sudden and long hop just
as it is going to be seized, and this is appreciated by the child as very
clever. The hunt, as a rule, does not result in much damage to the
grasshoppers, the ingenious escape being the greater part of the pleasure.
These _Stenobothri_ are remarkable for their variation in colour, and it is
thought by some that they frequent spots where they find themselves a match
with their surroundings. There is, however, little or no information of
importance on this point extant. _Mecostethus grossus_ (Fig. 173), though
larger, is very like the common field grasshoppers, but appears to have
become rare since the fens were drained. The two curious little
grasshoppers of the genus _Tettix_ (Fig. 179) are not uncommon. In addition
to these Acridiidae, three species of migratory locusts are occasionally
met with in Britain, viz. _Pachytylus cinerascens_ (Fig. 177), _P.
migratorius_, and _Schistocerca peregrina_ (Fig. 84); this latter we have
already alluded to as being probably the locust mentioned in the book of
Exodus.
Acridiidae have never been found in amber, owing possibly to their large
size and strength. There are but few fossil forms known, and these do not
extend farther back in time than the Mesozoic {309}epoch. Several forms,
including three peculiar genera, have been found in the Tertiary strata at
Florissant. The remains from the Mesozoic layers are apparently very
fragmentary and obscure.
Brongniart has instituted a family of Insects under the name
Palaeacrididae[242] for some fossil Insects from the Carboniferous strata
at Commentry. He considers that these Insects were abundant in the epoch of
the Carboniferous strata.
The very large number of genera and species of Acridiidae have been
recently arranged in nine tribes by Brunner von Wattenwyl:—
1. Feet without a claw-pad.[243] [Pronotum covering all the body.]
Tegmina lobe-like. Tribe 1. TETTIGIDES. (Figs. 179, 180, _Tettix_,
_Xerophyllum_, _Cladonotus_.)
1′. Feet with a claw-pad.
2. Antennae shorter than the anterior femora.
3. Head short, as if compressed from in front.
4. Body bladder-like, inflated.[244] [Pronotum covering half the
abdomen.] South African species. Tribe 2. PNEUMORIDES. (Fig. 182,
_Pneumora scutellaris_.)
4′. Body ordinary. Tribe 3. MASTACIDES. (Fig. 181, _Mastax
guttatus_.)
3′. Head very elongate. [Body apterous or sub-apterous.] Tribe 4.
PROSCOPIIDES. (Fig. 178, _Cephalocoema lineata_.)
2′. Antennae longer than the anterior femora.
3. Prosternum unarmed.
4. The plane of the vertex of the head meeting the plane of the
front of the head as an angle. The former produced or declivous.
The face looking down. Tribe 5. TRYXALIDES. (Fig. 165, _Tryxalis
nasuta_; Fig. 173, _Mecostethus grossus_.)
4′. Planes of the vertex and front of the head connected in a
rounded manner. Face looking forwards. Tribe 6. OEDIPODIDES.
(Fig. 177, _Pachytylus_; Fig. 185, _Methone_.)
3′. Prosternum with an elevated lamina in front, either irregularly
swollen or mucronate.
4. Foveoles of the vertex superior, contiguous, forming the apex of
the vertex. Face looking much downwards. Tribe 7. PYRGOMORPHIDES.
(Fig. 183, _Pyrgomorpha grylloides_.)
4′. Foveoles of the vertex, either superior (but not forming the
apex of the vertex), or lateral, or inferior, or quite obsolete.
{310}5. Foveoles superior, open behind. Prosternum irregularly
swollen, rarely mucronate. Tribe 8. PAMPHAGIDES. (Fig. 184,
_Xiphocera asina_.)
5′. Foveoles lateral or inferior, closed behind or (usually)
entirely obsolete. Prosternum distinctly mucronate or
tuberculate. Tribe 9. ACRIDIIDES. (Fig. 84, _Acridium
peregrinum_; Fig. 176, _Caloptenus spretus_.)
{311}CHAPTER XIII
ORTHOPTERA _CONTINUED_—LOCUSTIDAE, GREEN GRASSHOPPERS, KATYDIDS
FAM. VII. LOCUSTIDAE—GREEN GRASSHOPPERS.
_Orthoptera, with very long delicate antennae composed of many more than
thirty joints; hind legs longer than the others, thicker at the base.
Tarsi with four joints. Front tibiae usually provided with tympanal
organs placed below the knee; stridulating apparatus of males, when
present, situate on the basal part of the tegmina. Females usually with
an elongate exserted ovipositor, formed by the apposition of six pieces.
Wingless forms numerous._
[Illustration: FIG. 187.—_Cyrtophyllus crepitans_, male. West Indies.]
An unfortunate confusion has long existed as to the term Locustidae, and
has resulted in the application of the name to a group of Insects that
contains none of the locusts of ordinary language. Some entomologists
therefore use the term Phasgonuridea for this family, but the great
majority prefer the term Locustidae.
{312}[Illustration: FIG. 188.—Development of wings in _Platycleis grisea_:
A, B, C, D, E, consecutive stages; _p_, prothorax; _m_, mesothorax; _mt_,
metathorax; _t_, tegmen; _w_, wing; _ab′_, position of first abdominal
segment. In C, D, and E, _m_ points to the part by which the _m_, shown in
A and B, is concealed; in D and E only the positions of _mt_ are indicated.
(After Graber.)]
The Locustidae are, as a rule, more fragile Insects than the Acridiidae,
from which they can be readily distinguished by the characters we have
mentioned in our definition. According to Dufour, there are no air vesicles
connected with the tracheal system in this family; possibly to this it may
be due that none of the family undertake the long flights and migratory
wanderings that have made some of the Acridiidae so notorious. Very little
is known as to the life histories of the members of this extensive family
of Orthoptera. Graber, however, has given some particulars as to the
development of _Platycleis grisea_, and of one or two other species. He
recognises five instars, but his first is probably really the second, as he
did not observe the Insect in its youngest condition. Although his figures
are very poor, we reproduce them, as they give some idea of the mode of
growth of the wings, and of the correlative changes in the thoracic
segments. It will be seen that in the first three of these instars the alar
organs appear merely as prolongations of the sides of the posterior two
thoracic rings, and that in D a great change has occurred in the position
of these segments, so that the alar organs are free processes, the two
posterior thoracic rings being insignificant in size in comparison with the
now greatly developed prothorax. In E the tegmen is shown fully developed,
the positions of some {313}of the rings covered by it being indicated by
the letters _m_, _mt_, _ab′_. These changes are very similar to those we
have described in Acridiidae, the chief difference being the greater
development of the dependent wing-pads previous to the fourth instar.
[Illustration: FIG. 189.—Front of head of _Copiophora cornuta_, female.
Demerara.]
The ocelli in Locustidae are much more imperfect than they are in
Acridiidae, and are frequently rudimentary or nearly totally absent, or
there may be but one instead of three. They are, however, present in a
fairly well-developed state in some species, and this is the case with the
one whose face we portray in Fig. 189, where the anterior of the three
ocelli is quite conspicuous, the other two being placed one on each side of
the curious frontal cone near its base. The peculiar head ornament shown in
this figure exists in both sexes, and something similar occurs in a large
number of Conocephalides. We have not the slightest idea of its import.
Individuals of one or more species of this curious South American genus are
occasionally met with alive in gardens near London. They are, no doubt,
imported as eggs, for they are sometimes met with in the juvenile state,
but in what way they are introduced is not known.
The ovipositor frequently attains a great length in these Insects, so as to
exceed that of the body. It is used in different ways, some of the family
depositing their eggs in the earth, perhaps in vegetable matter under the
surface; but other species place the ova in twigs or stems of plants,
arranging them in a very neat and compact manner in two series, as depicted
by Riley[245] in the case of _Microcentrum retinerve_ (Fig. 190). These
eggs are laid in the autumn, and in the following spring become more
swollen before hatching. The Insect undergoes a moult during the process of
emerging from the egg. By the time the emergence is completed the
_Microcentrum_ has expanded so much {314}in size that it is a matter of
astonishment how it can ever have been packed in the egg; the young
commence jumping and eating leaves in a few minutes. Including the ecdysis
made on leaving the egg, they cast their skins five times. The
post-embryonic development occupies a period of about ten weeks. The larvae
eat their cast skins. When the final moult occurs the tegmina and wings are
at first quite soft and colourless, but within an hour they assume their
green colour. These Insects, as remarked by Riley, make interesting pets.
The people of the Amazon valley are in the habit of keeping a species in
cages, and our British _Locusta viridissima_ does very well in confinement.
One of the most curious habits of these Locustidae is a constant licking of
the front paws. Riley says that _M. retinerve_ bestows as much attention on
its long graceful antennæ as many a maiden does upon her abundant tresses,
the antennæ being drawn between the jaws and smoothed by the palpi. This
American naturalist also tells us that he reared three successive broods in
confinement, and that the Insects gradually deteriorated, so that the eggs
of the third generation failed to hatch.
[Illustration: FIG. 190.—Eggs of Katydid (_Microcentrum retinerve_): A, the
two series at deposition; B, side view of a single series. (After Riley.)]
The ovipositor, which is one of the most characteristic features of the
Locustidae, is not present in the newly-hatched Locustid (Fig. 191, A), the
organ being then represented only by two papillae placed on the penultimate
segment. The structure and development of the ovipositor in _Locusta
viridissima_ have been described by Dewitz.[246] Fig. 191, A, shows the
young Insect taken from the egg just as it is about to emerge. The abdomen
consists of ten segments, the terminal one bearing at its extremity two
processes, the cerci, _a′_. These persist throughout the life of the
Insect, and take no part in the formation of the ovipositor. The tenth
segment subsequently divides into two (_a_, _a'′_, Fig. 191, C), giving
rise to the appearance of eleven abdominal segments, and of the ovipositor
springing from the antepenultimate.
{315}[Illustration: FIG. 191.—Development of ovipositor of _Locusta
viridissima_: _a_, terminal segment; _a′_, cerci; _a″_, secondary division
of terminal segment; _b_, penultimate (ninth) segment; _b′_, primary
papillae of this segment; _b″_, secondary divisions thereof; _c_, eighth
segment; _c′_, its papillae. (After Dewitz.) A, embryo ready for emergence;
B, portion of integument of the ventral plates of eighth and ninth
segments; C, the appendages in a condition somewhat more advanced than they
are in A.]
[Illustration: FIG. 192.—Structure of ovipositor of _Locusta viridissima_:
A, arrangement of parts at base, _c′_ being separated and turned outwards;
B, transverse section. The parts of the appendage bear the same lettering
as in Fig. 191. (After Dewitz.)]
Near to one another, on the middle of the ventral aspect of the true ninth
abdominal segment, are seen the two papillae (_b′_), which at first are the
only visible indications of the future ovipositor. If, however, the
integument be taken off and carefully examined, it will be found that there
exist on the eighth abdominal plate two spots, where there is a slight
thickening and prominence of the integument (Fig. 191, B, _c_). From these
two spots the two lower rods of the ovipositor are produced; these two,
together with the two growths from the ninth segment, form the four
external rods of the ovipositor. Inside these there exist in the completed
structure two other rods (Fig. 192, B, _b'′_). These are produced by a
growth from the inner parts of the two papillae of the ninth segment. The
relations of the six rods in their early condition are shown in Fig. 191,
C, where the two primary papillae _b′_ of the ninth segment are seen with
their secondary offshoots _b″_; _c′_ being the papillae of the eighth
segment. The subsequent relations of the pieces are shown in Fig. 192; A
exhibiting the base of the organ with the lower rods turned on one side to
show the others, the shaded parts indicating {316}muscular attachments; B
is a transverse section of the organ. In these figures the different parts
of the appendages bear the same lettering as they do in Fig. 191. It will
be seen that in the completed structures the parts _c′_ have become very
intimately connected with the parts _b′_ and _b″_, which belong to another
segment.
The Locustidae resemble the Acridiidae in the possession of specialised
ears and sound-producing organs; neither of these is, however, situate in
the same part of the body as in Acridiidae. The ears of Locustidae are
placed on the front legs, below the knee; a tympanum (Fig. 193, A), or a
crack giving entrance to a cavity in which the tympanum is placed (Fig.
193, B), being seen on each side of each of the anterior pair of limbs. In
this family, as in the Acridiidae, three kinds of ear are recognised
according to the condition of the tympanum, which is either exposed (Fig.
193, A) or closed by an overgrowth of the integument (Fig. 193, B), or in a
condition to a certain extent different from either of these. The existence
of ears placed on the legs is a curious fact, but it is beyond doubt in the
Locustidae, and there is good reason for believing that analogous organs
exist in this situation in other Insects that have special means of
sound-production, such as the ants and the Termites.
[Illustration: FIG. 193.—Ears of Locustidae: A, portion of front leg of
_Odontura serricauda_, adult; _p_, prominence of integument; _r_, rim of
ear; _T_, tympanum; _b_, thickened area thereof; _Fu_, remains of groove in
which the structure was developed. B, portion of front leg of _Thamnotrizon
apterus_; _i_, inner margin; _a_, slit-like external aperture of ear; _di_,
overlapping cover of the ear. (After Graber.)]
The structure of these organs in the Locustidae has been investigated by
Graber,[247] and their acoustic functions placed beyond doubt, though to
what special kind of sounds they may be sensitive is not ascertained, this
point being surrounded by even greater difficulties than those we have
discussed in the case of the Acridiidae. In the Locustidae there is a
special structure of a remarkable nature in connexion with the ears. In
Acridiidae {317}a stigma is placed close to the ear, and supplies the
internal structures of the organ with air. There are no stigmata on the
legs of Insects, consequently admission of air to the acoustic apparatus in
Locustidae is effected by means of a gaping orifice at the back of the
prothorax, just over the base of the front leg (Fig. 101); this
communicates with its fellow of the other side, and from them there extend
processes along the femora into the tibiae, where they undergo dilatation,
so as to form vesicular cavities, one of which is in proximity to each drum
of the ear. These leg-tracheae are not connected with the ordinary tracheal
system; the prothoracic stigma exists in close proximity to the acoustic
orifice we have described, but is much smaller than it. It is not yet clear
why the acoustic apparatus should require a supply of air apart from that
which could be afforded by the ordinary tracheal system. This special
arrangement—to which there is hardly a parallel in Insect anatomy—has still
to be accounted for; we do not know whether the necessity for it may be
connected with the respiratory system or the acoustic organ.
[Illustration: FIG. 194.—Diagram of arrangement of parts of the ear as seen
in transverse section of the tibia of a Locustid. A, J, V, H, outer, inner,
anterior, posterior aspects of leg; _a_, _d_, thin part of integument
forming anterior tympanum; _b_, _c_, thicker portion of same; _f_, _g_,
posterior tympanum; _a_, _f_, and _d_, _h_, _g_, thick portions of
integument; _i_, _k_, internal protuberances of same; _l_, _m_, _n_, _o_,
walls of the anterior tracheal vesicle, _vTr_; _p_, _q_, _s_, _r_, walls of
the posterior tracheal vesicle, _hTr_; _o″_, projection of tympanal orifice
of prothorax; _tr-n_, tracheal nerve-end organ, _crista acustica_; _st_,
rod; _de_, curtain-membrane; _hn_, _e_, supra-tympanal, nerve-end organ;
_hn_, ganglion cells; _st′_, rods; _e_, point of integumental fixation of
nerve endings. (After Graber.)]
The chief features of the acoustic apparatus of the legs of Locustidae will
be gathered from the accompanying diagrammatic transverse section through
the tibia. In this figure the deep black parts indicate the outer wall of
the tibia and its prolongations, the white spaces indicate the parts filled
with air, while the dotted portions are occupied by blood or some of the
body organs;[248] the {318}circular space _o″_ is not part of the actual
structure, but represents the area of the external acoustic orifice of the
prothorax; it is not, however, so large as it should be.
Although the tibial ears of Locustidae are very perfect organs, there is
great difficulty in deciding on the exact nature of their functions. They
would appear to be admirably adapted to determine the precise locality from
which a sound proceeds, especially in those cases—and they are the highest
forms—in which the tympanum is placed in a cavity the external orifice of
which is a slit (Fig. 193, B); for the legs can be moved in the freest
manner in every direction, so as to bring the drum into the most direct
line of the vibrations. But as to what kinds of vibrations may be
perceived, and the manner in which they may be transmitted to the nerves,
there is but little evidence. On reference to the diagram it will be
noticed that the tympanum, the tympanal vesicles, and the nervous apparatus
are not in close connexion, so that even the mode by which the impulses are
transmitted is obscure.
The musical organs of the Locustidae are different from those of the
Acridiidae, and are invariably situate on the basal part of the tegmina.
They are found, in the great majority of cases, only in the male; in the
tribes Ephippigerides and Callimenides they exist in each sex. One of the
wings bears a file on its inner surface, while the other—on the right side
of the body—is provided with a sharp edge placed on a prominent part of its
inner margin. By slightly tilting the tegmina and vibrating them rapidly,
the edge passes under the file, and a musical sound is produced. These
structures are limited to the small anal area of the wing, and when the
tegmina are very greatly reduced in size, it is this part that still
remains. There is much variety in the details of the structure. The
nervures of this part of the tegmina are different in the male from what
they are in the female, and, moreover, the two wing-covers of the male
differ from one another. It is apparently the vibrations of the right
tegmen that produce the sound, and this part usually bears a space of a
glassy nature, which probably improves the character of the sound produced.
Our chief British songster of this group, _Locusta viridissima_, is only
provided with phonetic organs (Fig. 195) of a somewhat imperfect character,
but in the genus _Mecopoda_ there is great perfection of the structures.
The anal areas of the two tegmina are in this case {319}very different;
that of the left one, which bears the file, being similar in texture to the
rest of the wing-cover, while the corresponding part of the other tegmen is
rigid and transparent, and greatly distorted, so as to create a cavity
which, no doubt, improves the sound; the scraper too is very perfectly
formed. The difference between this form of musical organ and that of _L.
viridissima_ is curious, inasmuch as in the better instrument the important
modifications are confined to one tegmen, while in the other form both
tegmina are largely changed. The difference appears to be that in _Locusta_
the left tegmen, as well as the right one, acts as a sounding-board, while
in _Mecopoda_ it does not do so, but when the wings are closed quite covers
and conceals the musical instrument.
The Locustidae, notwithstanding the fact that their alar organs are
generally more ample than those of the Acridiidae, seem to be, as a rule,
of more sedentary habits, and more nocturnal in their activity. The musical
powers of the different species are very varied. _Locusta viridissima_
produces a shrill and monotonous but not disagreeable, sound, and is
capable of sustaining it for a quarter of an hour without any intermission,
except a break for the sake of starting again immediately with greater
force, like a performer on a flute. It occasionally chirps in the day, but
the act is then very brief. Bates informs us that one of these singing
grasshoppers, called Tananá by the natives of the Amazon valley, is much
admired for its singing, and is kept in little cages. The Amazonian
naturalist thought the music of this species superior to that of any other
Orthopterous Insect he had heard. The name of this grasshopper is
_Thliboscelus camellifolius_. It is very similar in appearance to
_Cyrtophyllus crepitans_, the Insect we have represented in Fig. 187.
[Illustration: FIG. 195.—Inner face of base of tegmina of _Locusta
viridissima_: A, the two wing-covers separated; B, in natural position with
mesonotum connecting them, showing file and edge scraping it; _a_, the
stridulating file; _b_, the rudimentary file on other tegmen.]
The most notorious of the musical Locustids are the Katydids {320}of North
America. There are several species of them—they belong, indeed, to more
than one genus,—but it seems that sounds somewhat resembling the words
Katy-did are perceptible in most of their performances. These sounds are
frequently repeated with slight variations—Katy-did, O-she-did,
Katy-did-she-did. Riley describes the music of the Katydid we represent in
Fig. 196 as follows:[249] "The first notes from this Katydid are heard
about the middle of July, and the species is in full song by the first of
August. The wing-covers are partially opened by a sudden jerk, and the
notes produced by the gradual closing of the same. The song consists of a
series of from twenty-five to thirty raspings, as of a stiff quill drawn
across a coarse file. There are about five of these raspings or trills per
second, all alike, and with equal intervals, except the last two or three,
which, with the closing of the wing-covers, run into each other. The whole
strongly recalls the slow turning of a child's wooden rattle, ending by a
sudden jerk of the same; and this prolonged rattling, which is peculiar to
the male, is invariably and instantly answered by a single sharp 'chirp' or
'tschick' from one or more females, who produce the sound by a sudden
upward jerk of the wings."
[Illustration: FIG. 196.—Katydid, _Microcentrum retinerve_. N. America.
(After Riley.)]
Pertinacity is one of the most curious features of the performance of
musical Locustids. One would say they desire to distinguish themselves as
much as possible. Harris says that _Cyrtophyllus concavus_ mounts on the
uppermost twigs of trees and there performs its Katy-did-she-did in rivalry
with others. He says even the female in this species gives forth a feeble
noise. Scudder says that some of the Katydids sing both by day and night,
but their day song differs from that of the night. "On a summer's day it is
curious to observe these little creatures suddenly {321}changing from the
day to the night song at the mere passing of a cloud, and returning to the
old note when the sky is clear. By imitating the two songs in the daytime
the grasshoppers can be made to respond to either at will; at night they
have but one note."
Although but little is known as to the habits of Locustidae, it is
ascertained that they are less exclusively herbivorous in their food habits
than the Acridiidae are; many seem to prefer a mixed diet. _Locusta
viridissima_ will eat various leaves and fruits, besides small quantities
of flesh. It has been recorded that a specimen in confinement mastered a
humble-bee, extracted with its mandibles the honey-bag, and ate this
dainty, leaving the other parts of the bee untouched. Many of the
Locustidae are believed to be entirely carnivorous. Brunner considers a
minority to be exclusively phytophagous. The species very rarely increase
to large numbers; this, however, occurs sometimes with _Orphania
denticauda_ and _Barbitistes yersini_ in Europe, and _Anabrus purpurascens_
in North America. We have already mentioned that the eggs of some species
are deposited in parts of plants, and of others in the earth. The British
_Meconema varium_ deposits its eggs in the galls of _Cynips_ in the autumn;
these eggs do not hatch till the following spring. _Xiphidium ensiferum_
has somewhat similar habits in North America, the gall selected for the
reception of the eggs being the scales formed by a species of _Cecidomyia_
on the leaves of willows. It has been ascertained that the development of
the embryo in the last-named species is commenced in the autumn, but is
suspended during the winter, being only completed in the following spring,
eight or nine months afterwards. We owe to Wheeler[250] a memoir on the
embryology of this Insect.
Some of the species have the peculiar habit of dwelling in caves. This is
especially the case with the members of the tribe Stenopelmatides (Fig.
197), which frequently possess enormously long antennae and legs, and are
destitute of alar organs and ears. The species with this habit, though
found in the most widely separated parts of the world, have a great general
resemblance, so that one would almost suppose the specimens found in the
caves of Austria, in the Mammoth cave of Kentucky, and in the rock-cavities
of New Zealand to be one {322}species, although they are now referred by
entomologists to different genera.
[Illustration: FIG. 197.—_Dolichopoda palpata_, male. Dalmatia. (After
Brunner.)]
[Illustration: FIG. 198.—Leaf-like tegmen of _Pterochroza ocellata_: _a_,
_a_, _a_, marks like those made by Insects on leaves.]
The Locustidae display in the greatest possible perfection that resemblance
of the tegmina to leaves which we mentioned when speaking of the general
characters of the Orthoptera. The wing-covers are very leaf-like in colour
and appearance in many Locustidae, but it is in the tribe Pseudophyllides
and in the South American genus _Pterochroza_ (Fig. 198) that the
phenomenon is most remarkable. The tegmina in the species of this genus
look exactly like leaves in certain stages of ripeness or decay. In the
tegmina of some of the species not only are the colours of faded leaves
exactly reproduced, but spots are present like those on leaves due to
cryptogamic growths. Perhaps the most remarkable feature of these
resemblances is the one pointed out by Brunner von Wattenwyl,[251] viz.
that the tracks and spots formed on leaves by the mining of Insects in
their tissues are also represented in the leaf-like wing-covers of the
_Pterochroza_; transparent spots (_a_, _a_, Fig. 198) being present, just
as they are in many leaves that have been attacked by Insects. Brunner was
so much impressed by these facts that he came to the conclusion that they
cannot be accounted for on the grounds of mere utility, {323}and proposed
the term Hypertely to express the idea that in these cases the bounds of
the useful are transcended. We will mention here another peculiar case of
resemblance described by Brunner as occurring in a Locustid. Two specimens
of a little Phaneropterid were brought from the Soudan by the Antinori
expedition, and have been described by Brunner under the name of
_Myrmecophana fallax_. The Insect is said to bear an extraordinary
resemblance to an ant. The most peculiar feature in the resemblance is
shown in Fig. 199, A, B. The most characteristic point in the external form
of an ant is the stalked abdomen, this structure being at the same time
quite foreign to the Orthoptera. In the other parts of the body and in the
colour generally, the _Myrmecophana_ resembles an ant, but the abdomen of
the Orthopteron is not stalked; it has, however, the appearance of being
so, in consequence of certain parts being of a white colour, as shown in
our figure. If abstraction be made of the white parts, the form of the
stalked abdomen of the ant is nicely reproduced. The specimens brought from
the Soudan were wingless and destitute of ovipositor, and may be immature,
but Brunner suggests that they may prove to be really mature, the
ovipositor, tegmina, and wings being permanently absent. The existence of a
long ovipositor would certainly detract greatly from the ant-like
appearance of the Orthopteron.
[Illustration: FIG. 199.—_Myrmecophana fallax._]
It is certain that the plant-like appearance of some of the Locustidae
renders them inconspicuous to the human eye in the situations they
frequent. It is a matter of common observation that though the noise of
their chirpings may be heard to such an extent as to make it certain that
many individuals must be in the immediate neighbourhood, yet at the same
time it may be most difficult to detect even a single individual. M. Boutan
noticed this phenomenon in the case of _Ephippigera rugosicollis_, and
tells us that the human eye can, with a little practice, acquire the art of
detecting these concealed creatures. This consists {324}apparently in
making use, not of a general inspection, but of a scrutiny of the outlines
of the leaves and twigs of a tree. By this means, when the eye is
accustomed to the task, the Insects can be detected with comparative ease;
much in the same way, M. Boutan says, as a figure, placed in an engraving
in such a way as to elude the eye, is appreciated with ease after the eye
has once perceived it.
Some of the Locustidae are provided with means of defence of a positive
nature. The Algerian _Eugaster guyoni_ ejects two jets of a caustic
orange-coloured fluid from two pores situate on the sides of the
mesosternum, and covered by the anterior coxae. This species is carnivorous
as well as herbivorous, and produces a sound more like humming than
stridulation.[252]
[Illustration: FIG. 200.—_Phasmodes ranatriformis_,female. Australia.
(After Westwood.)]
We have previously pointed out that some of the Acridiidae resemble the
stick-Insects rather than the members of their own group; and similar cases
occur amongst the Locustidae. Such a resemblance has, however, only been
found in a few species of the tribe Prochilides. We figure one of these,
_Phasmodes ranatriformis_, a native of South-West Australia. The very
elongate linear form and the total absence of alar organs give this Insect
a considerable resemblance to the stick-Insects or apterous Phasmidae.
_Prochilus australis_ is allied to this curious Locustid, but the alar
organs are present in both sexes, and the Insect bears a great resemblance
to the winged Phasmidae. This is due not only to the general form and
colour, but also to the fact that the tegmina are very narrow, which
{325}causes them to look like the coloured slip on the anterior parts of
the wings of some of the Phasmidae (cf. p. 266). Another case of a Locustid
with elongate, slender form is found in the extraordinary _Peringueyella
jocosa_ of South Africa, a member of the tribe Sagides. It has minute
organs of flight, and reproduces, to a considerable extent, the form and
appearance of Proscopides or of some Tryxalides.[253]
[Illustration: FIG. 201.—_Schizodactylus monstrosus_, male. Natural size.
East India.]
We follow Brunner in placing among the Locustidae the large Insect we
represent in Fig. 201. It is remarkable on account of its tegmina and
wings; these have their extremities much prolonged and curled; moreover,
the flat interior area and the abruptly {326}deflexed exterior area make
them look more like the wings of Gryllidae. This species has no ocelli, and
is said to be destitute of ears. The inflated condition of the anterior and
middle tibiae suggest that it possesses auditory structures, though there
appears to be no external opening for them. This Insect is found in India,
where it is said to be common on the banks of sandy rivers, living there in
burrows of the depth of three feet. Very little is known, however, as to
this curious Insect. It has recently been reported[254] as being injurious
to tobacco and other crops on high ground in Durbungha by cutting off their
roots. The local name for the Insect is _bherwa_. We should think it
somewhat doubtful whether this refers really to _S. monstrosus_.
[Illustration: FIG. 202.—_Anostostoma australasiae_, male. Australia.]
In number of species the Locustidae are perhaps scarcely inferior to the
Acridiidae, and in variety of form they surpass this latter family. Many of
the most gigantic forms are apterous, and these very often have a repellant
aspect. The genus _Anostostoma_ is remarkable for its large head. Allied to
it is _Deinacrida heteracantha_, the "Weta-punga" of the New Zealand
natives, an Insect formerly abundant in the forests north of Auckland, but
of late years become extremely rare. The head and body of this Insect may
measure more than 2½ inches in length, and when the antennae and legs are
stretched out the total length may be 14 or 15 inches. Although bulky and
absolutely wingless, yet, as Buller informs us,[255] it climbs with
agility, and is sometimes found on the topmost branches of lofty trees.
When disturbed it produces a clicking, accompanied by a slow movement
{327}of its hind legs. A second species, _D. thoracica_, lives in decayed
wood, and a third, _D. megacephala_, is remarkable from the very large size
of the head and mandibles in the male sex. The fact that a clicking noise
is produced by the Weta-punga is of some interest, for the genus
_Deinacrida_ is among the Locustidae that possess ears, but are said to be
destitute of sound-producing organs.
Amongst the most remarkable of the Locustidae are the two species of which
Brongniart has recently formed the genus _Eumegalodon_ and the tribe
Eumegalodonidae, which is not included in Brunner's table of the tribes of
Locustidae. The ovipositor is large and sabre-shaped; the male is unknown.
The genus _Megalodon_ is placed by Brunner in the tribe Conocephalides; it
also consists of extremely remarkable Insects.
[Illustration: FIG. 203.—_Eumegalodon blanchardi_, female. Borneo. × ⅘.
(After Brongniart.)]
The Locustidae appear to be of slow growth, and the autumns of Britain are
usually not warm enough for them. Hence we have but nine British species,
and of this number only three or four are known to occur north of the
Thames. The only one that attracts attention is _Locusta viridissima_,
which in some districts of the south of England occurs in considerable
numbers, and attests its presence by its peculiar music. It is called the
green grasshopper.
{328}The geological record is rather obscure in the matter of Locustidae.
Scudder considers that a fair number of Tertiary forms are known, and says
that they represent several of the existing tribes and genera. One or two
have been found in Mesozoic rocks.
TABLE OF THE TRIBES OF LOCUSTIDAE
1. Tarsi more or less depressed.
2. Front tibiae furnished with auditory cavities.
3. Antennae less distant from the summit of the occiput than from the
labrum; inserted between the eyes.[256]
4. First two joints of the tarsi laterally smooth. (Posterior
tibiae furnished on each side with an apical spine.) Tribe 1.
PHANEROPTERIDES. (Fig. 196, _Microcentrum_; Fig. 199,
_Myrmecophana_. Fig. 101, _Poecilimon affinis_.)
4′. First two joints of the tarsi laterally, longitudinally
sulcate.
5. Foramina of the anterior tibiae normally open. (Fig. 193, A.)
6. Posterior tibiae furnished on each side with apical spines.
7. Prosternum unarmed. Tribe 2. MECONEMIDES.
7′. Prosternum bispinose or bituberculate. Tribe 3.
MECOPODIDES.
6′. Posterior tibiae with no apical spines. (Head prognathous.)
Tribe 4. PROCHILIDES. (Fig. 200, _Phasmodes_.)
5′. Foramina of the anterior tibiae forming a chink, or protected
by a scale. (Fig. 193, B.)
6. Anterior tibiae with no apical spines.
7. Margins of the scrobes[257] of the antennae prominent.
Tribe 5. PSEUDOPHYLLIDES. (Fig. 187, _Cyrtophyllus
crepitans_; Fig. 198, _Pterochroza ocellata_.)
7′. Margins of the scrobes of the antennae not prominent.
8. Posterior tibiae furnished above on each side with
apical spines, or with a single spine on the side.
9. Posterior tibiae either furnished with apical spines
on each side, or only on the inner side. Tribe 6.
CONOCEPHALIDES. (Fig. 189, _Copiophora cornuta_.)
9′. Posterior tibiae furnished above with an apical spine
placed only on the outer side. Tribe 7. TYMPANOPHORIDES.
8′. Posterior tibiae without apical spines. Tribe 8.
SAGIDES.
6′. Anterior tibiae furnished with an apical spine on the inner
side.[258]
{329}7. The first joint of the posterior tarsi destitute of a
free sole-lobe. Tribe 9. LOCUSTIDES.
7′. The first joint of the posterior tarsi furnished with a
free sole-lobe. Tribe 10. DECTICIDES.
3′. Antennae more distant from the summit of the occiput than from
the labrum, inserted either beneath the eyes or on their inferior
border. Tegmina and wings greatly abbreviate, scale-like; when
tegmina are present they are furnished in each sex with a tympanum.
4. Third joint of the posterior tarsi shorter than the second. Both
anterior and posterior tibiae furnished on each side with a spine.
Tribe 11. CALLIMENIDES.
4′. Third joint of posterior tarsi longer than the second joint.
Anterior tibiae with no apical spine on the inner side, and
posterior tibiae with no apical spine on the outer side.
5. Antennae inserted at the edge of the eyes. Pronotum unarmed.
Tegmina present in each sex. Anterior tibiae furnished on the
outer side with an apical spine. Posterior tibiae furnished
beneath with four apical spines. Tribe 12. EPHIPPIGERIDES.
5′. Antennae inserted distinctly below the eyes. Pronotum
spinous. Elytra in the females wanting. Anterior tibiae without
apical spine on either side. Posterior tibiae beneath with two
apical spines or with none. Tribe 13. HETRODIDES.
2′. Anterior tibiae without auditory cavities. Tegmina with no
tympanum. Tribe 14. GRYLLACRIDES. (Fig. 201, _Schizodactylus
monstrosus_.)
1′. Tarsi distinctly compressed (most of the species apterous.) Tribe 15.
STENOPELMATIDES. (Fig. 202, _Anostostoma australasiae_; Fig. 197,
_Dolichopoda palpata_.)
{330}CHAPTER XIV
ORTHOPTERA _CONTINUED_—GRYLLIDAE, CRICKETS
FAM. VIII. GRYLLIDAE—CRICKETS.
_Antennae very slender, generally long and setaceous; hind legs long,
saltatorial. Tegmina with the outer portion deflexed on to the side of
the body, and with the inner part lying flat on the body. Tarsi usually
three-jointed (rarely two- or four-jointed). Female with a long
ovipositor (except in Gryllotalpides). Apterous forms numerous._
The Gryllidae are closely connected with the Locustidae, the musical and
auditory organs being in both similarly situate, and the female in both
possessing, in most of the tribes, an elongate exserted ovipositor. The two
families differ in the number of joints of the tarsi, in the form of the
tegmina, and in the fact that in Gryllidae the portion of the wing modified
for musical purposes consists of a larger portion of the organ—according to
de Saussure, the discoidal as well as the anal area.
[Illustration: FIG. 204.—House-cricket, _Gryllus_ (_Acheta_) _domesticus_,
male.]
The family would be a very natural one if we were to exclude from it the
mole-crickets which have fossorial front legs and no ovipositor, and the
Tridactylides, which also are {331}destitute of ovipositor, and have short
antennae, consisting of about ten joints.
The head is generally very large; ocelli are present, though usually
imperfect; the extremity of the body bears a pair of remarkably long cerci.
The hind tibiae are usually armed with very strong spines; the first joint
of the hind tarsus is elongate, and terminates in two spines, between which
the small second joint is often almost completely concealed; the feet are
not provided beneath with pads, but only bear remote setae.
The alar organs are difficult of comprehension, and different opinions
prevail as to their morphology. The tegmina are extremely different to the
hind wings, and never attain large dimensions, neither do they exhibit any
leaf-like or ornamental structures. In the genus _Pteroplistus_ they are
formed somewhat like the elytra of Coleoptera, and close over the back of
the Insect in a fashion very like that found in beetles. According to
Brunner the larger part of the tegmen—which, as we have said, reposes flat
on the back of the Insect—represents merely the anal area, and all the
other parts must be sought in the smaller, deflexed portion of the
wing-cover. De Saussure's opinion, to a somewhat different effect, we have
already mentioned. The tegmina of the male are extremely different from
those of the female, so that it is a matter of much difficulty to decide
what nervures correspond.[259]
[Illustration: FIG. 205.—Tegmina (sinistral) of the house-cricket. A, male,
inner aspect; B, female, outer aspect: _a_, inner margin; _b_, outer
margin; _c_, nervure bearing stridulating file.]
The wing-covers of the male differ from those of the Locustidae, inasmuch
as the pair are of similar formation, each bearing a stridulating file on
its lower aspect. This file projects somewhat inwards, so that its position
is marked on the outer aspect of the wing-cover by a depression. Usually
the right tegmen overlaps the other, an arrangement contrary to that which
prevails in other Orthoptera. The wings are ample and delicate; they
possess numerous nervures that are not much forked and have a {332}simple,
somewhat fan-like arrangement; the little transverse nervules exhibit only
slight variety. These wings are frequently rolled up at the apex, and
project beyond the body like an additional pair of cerci (Fig. 204). The
abdomen is chiefly remarkable for the large development of the pleura, the
stigmata being consequently very conspicuous. The cerci are not jointed,
though they are flexible and, often, very long; they bear a variety of
sense-organs (Fig. 67). The saltatorial powers of the crickets are
frequently considerable.
Graber has observed the post-embryonic development of the field-cricket,
_Gryllus campestris_, though unfortunately not from the very commencement,
so that we do not know whether there are five, six, or seven ecdyses; the
number is probably either six or seven. The manner in which the alar organs
are developed is similar to that we have described and figured in the
Locustidae. In the earlier instars there is a slight prolongation of each
side of the meso- and meta-notum, but about the middle of the development a
considerable change occurs—the rudimentary organs then become free
appendages and assume a different position.
The Gryllidae possess a pair of tympana on each front leg, but these organs
contrast with those of the Locustidae in that the pair on each leg usually
differ from one another, the one on the outer or posterior aspect being
larger than that on the inner or front face of the leg.
The ears of the Gryllidae have not been so well investigated as those of
the Locustidae, but are apparently of a much less perfect nature. No
orifice for the admission of air other than that of the prothoracic stigma
has been detected, except in _Gryllotalpa_. On the other hand, it is
said[260] that in addition to the tibial organs another pair of tympana
exists, and is seated on the second abdominal segment in a position
analogous to that occupied by the ear on the first segment of Acridiidae.
The musical powers of the crickets are remarkable, and are familiar to all
in Europe, as the performance of the house-cricket gives a fair idea of
them. Some of the Insects of the family are able to make a very piercing
noise, the note of _Brachytrypes megacephalus_ having been heard, it is
said, at a distance of a mile from where it was being produced. The mode of
{333}production is the same as in the Locustidae, rapid vibration of the
tegmina causing the edge of one of them to act on the file of the other.
The mole-cricket, _Gryllotalpa vulgaris_—the _Werre_ of the Germans,
_Courtilière_ of the French—is placed with a few allies in a special group,
Gryllotalpides, characterised by the dilated front legs, which are
admirably adapted for working underground. Like the mole, this Insect has a
subterranean existence. It travels in burrows of its own formation, and it
also forms beneath the surface a habitation for its eggs and family. Its
habits have been alluded to by Gilbert White,[261] who tells us that "a
gardener at a house where I was on a visit, happening to be mowing, on the
6th of May, by the side of a canal, his scythe struck too deep, pared off a
large piece of turf, and laid open to view a curious scene of domestic
economy: there were many caverns and winding passages leading to a kind of
chamber, neatly smoothed and rounded, and about the size of a moderate
snuff-box. Within this secret nursery were deposited near a hundred eggs of
a dirty yellow colour, and enveloped in a tough skin, but too lately
excluded to contain any rudiments of young, being full of a viscous
substance. The eggs lay but shallow, and within the influence of the sun,
just under a little heap of fresh moved mould like that which is raised by
ants."
[Illustration: FIG. 206.—Front leg of the mole-cricket. A, outer; B, inner
aspect: _e_, ear-slit.]
The front legs are remarkable structures (Fig. 206), being beautifully
adapted for burrowing; the tibiae and tarsi are arranged so as to act as
shears when it may be necessary to sever a root. The shear-like action of
the tarsus and tibia is very remarkable; the first and second joints of the
former are furnished with hard processes, which, when the tarsus is moved,
pass over the edges of the tibial teeth in such a way as to be more
effective than a pair of shears. In consequence of its habit of cutting
roots, {334}the mole-cricket causes some damage where it is abundant. It is
now a rare Insect in England, and is almost confined to the southern
counties, but in the gardens of Central and Southern Europe it is very
abundant. Its French name _courtilière_ is supposed to be a corruption of
the Latin _curtilla_. Its fondness for the neighbourhood of water is well
known. De Saussure says that in order to secure specimens it is only
necessary to throw water on the paths between the flower-beds of gardens
and to cover the wetted places with pieces of board; in the morning some of
these Insects are almost sure to be found under the boards disporting
themselves in the mud. The Gryllotalpae swim admirably by aid of their
broad front legs.
Ears exist in the mole-cricket, and are situate on the front leg below the
knee, as in other Gryllidae, although it seems strange that a leg so
profoundly modified for digging and excavating as is that of the
mole-cricket should be provided with an ear. In _Gryllotalpa_ the ear is
concealed and protected by being placed in a deep slit or fold of the
surface, and this depression is all that can be seen by examination of the
exterior (Fig. 206, _e_). In the allied genus _Scapteriscus_ the tympanal
membrane is, however, destitute of special protection, being completely
exposed on the surface of the leg.
Although the tegmina or upper wings in _Gryllotalpa_ are of small size, yet
the true wings are much more ample; they are of delicate texture and
traversed by many nearly straight radii, so that they close up in the most
complete manner, and form the two long delicate, flexible processes that in
the state of repose may be seen projecting not only beyond the tegmina, but
actually surpassing the extremity of the body hanging down behind it, and
looking like a second pair of cerci.
The mole-cricket is believed to be chiefly carnivorous in its diet, though,
like many other Orthoptera, it can accommodate its appetite to parts of the
vegetable as well as of the animal kingdom. The Insect is capable of
emitting a sound consisting of a dull jarring note, somewhat like that of
the goat-sucker. For this purpose the tegmina of the males are provided
with an apparatus of the nature we have already described, but which is
very much smaller and less elaborate than it is in the true crickets.
{335}[Illustration: FIG. 207.—Alimentary canal and appendages of the
mole-cricket: _a_, head; _b_, salivary glands and receptacle; _c_, lateral
pouch; _d_, stomato-gastric nerves; _e_, anterior lobes of stomach; _f_,
peculiar organ; _g_, neck of stomach; _h_, plicate portion of same; _i_,
rectum; _k_, lobulate gland; _l_, extremity of body; _m_, Malpighian tubes.
(After Dufour.)]
The alimentary canal and digestive system of _Gryllotalpa_ present
peculiarities worthy of notice. Salivary glands and reservoirs are present;
the oesophagus is elongate, and has on one side a peculiar large pouch
(Fig. 207, _c_); beyond this is the gizzard, which is embraced by two lobes
of the stomach. This latter organ is, beyond the lobes, continued backwards
as a neck, which subsequently becomes larger and rugose-plicate. On the
neck of the stomach there is a pair of branching organs, which Dufour
considered to be peculiar to the mole-cricket, and compared to a spleen or
pancreas. The single tube into which the Malpighian tubules open is seated
near the commencement of the small intestine. These tubules are very fine,
and are about one hundred in number. The arrangement by which the
Malpighian tubules open into a common duct instead of into the intestine
itself appears to be characteristic of the Gryllidae, but is said to occur
also in _Ephippigera_, a genus of Locustidae. According to Leydig[262] and
Schindler the Malpighian tubules are of two kinds, differing in colour,
and, according to Leydig, in contents and histological structure. Near the
posterior extremity of the rectum there is a lobulated gland having a
reservoir connected with it; this is the chief source of the foetid
secretion the mole-cricket emits when seized. The nervous chain consists of
three thoracic and four abdominal ganglia; these latter do not extend to
the extremity of the body; {336}the three anterior of the four ganglia are
but small, the terminal one being much larger.
The number of eggs deposited by a female mole-cricket is large, varying, it
is said, from 200 to 400. The mother watches over them carefully, and when
they are hatched, which occurs in a period of from three to four weeks
after their deposition, she supplies the young with food till their first
moult; after this occurs they disperse, and begin to form burrows for
themselves.
It has been said that the young are devoured by their parents, and some
writers have gone so far as to say that 90 per cent of the progeny are thus
disposed of. M. Decaux, who has paid considerable attention to the economy
of the mole-cricket,[263] acquits the mother of such an offence, but admits
that the male commits it. The number of eggs in one nest is said to be
about 300.
The embryonic development of the mole-cricket has been studied by
Dohrn[264] and Korotneff,[265] and is considered by the former to be of
great interest. The tracheae connected with each stigma remain isolated,
while, according to Korotneff, the development of the alimentary canal is
not completed when the young mole-cricket is hatched. Perhaps it may be
this condition of the digestive organs that necessitates the unusual care
the mother bestows on her young.
[Illustration: FIG. 208.—_Cylindrodes kochi._ Australia. A, outline of the
Insect with five of the legs and the extremity of the body mutilated; B,
middle leg. (After de Saussure.)]
The genus _Cylindrodes_ (Fig. 208, _C. kochi_) comprises some curious and
rare Insects of elongate, slender form. They are natives of Australia,
where the first species known of the genus was found in Melville Island by
Major Campbell, from whom we learn that these Insects burrow in the stems
of plants, and are so destructive that he was unable to keep a single plant
in his greenhouse on account of the ravages of _Cylindrodes campbellii_.
The form of these Insects is beautifully adapted to {337}their habits, the
body being contracted in the middle in such a way as to permit the middle
and hind legs to be packed against it, so that the cylindrical form is not
interfered with by these appendages while the excavating anterior legs are
at work in front of the Insect. The abdomen has nine segments; the terminal
one, said to be remarkably long and destitute of cerci, is not shown in our
figure.
The genus _Tridactylus_ is considered by de Saussure to form, with its ally
_Rhipipteryx_, a division of Gryllotalpinae, but they are treated, perhaps
more correctly, by Brunner as a separate tribe. _T. variegatus_ (Fig. 209)
is a small Insect, abundant in sandy places on the banks of rivers in
Southern Europe,—extending on the Rhone as far north as Geneva,—and is
remarkable for its great power of leaping, and for the rapidity with which
it can burrow in the sand. This anomalous Insect has only ten joints to the
antennae. Its alar organs are imperfect, and not like those of other
Gryllidae in either form or neuration. The hind legs are of peculiar
structure, the tibiae terminating in two processes between which is situate
a rudimentary tarsus. Near the extremity of the tibia there are some
plates, forming two series, that can be adpressed to the tibia, or extended
as shown in our figure. The body is terminated by four rather short, very
mobile processes; the upper pair of these are each two-jointed, and are
thought by de Saussure and Haase[266] to be cerci; the inferior pair, being
articulated processes of the anal segment, their presence in addition to
cerci is remarkable. It is difficult to distinguish the sexes of this
Insect.
[Illustration: FIG. 209.—_Tridactylus variegatus_, France.]
The exotic genus _Rhipipteryx_ is allied to _Tridactylus_. It is widely
distributed in South America, but the little Insects that {338}compose it
are rare in collections, their saltatorial powers no doubt making it
difficult to catch them; little is known as to their habits. In the
undescribed Amazonian species we figure (Fig. 210), the wings, instead of
being mere rudiments, as in _Tridactylus_, are elongate and project beyond
the body; they are of a blue-black colour, and arranged so as to look as if
they were the abdomen of the Insect; they, moreover, have a transverse
pallid mark, giving rise to an appearance of division. It is difficult to
form any surmise as to the nature of so curious a modification of the
wings.
The Tridactylides have no tympana on the legs, and their affinity with the
Gryllidae is very doubtful. Dufour thought _T. variegatus_ to be more
allied to the Acridiidae. He based this opinion chiefly on some points of
the internal anatomy, but pointed out that _Tridactylus_ differs from the
Acridiidae in having no air-sacs in the body.
[Illustration: FIG. 210.—_Rhipipteryx_ sp., Amazon valley.]
Not many of the Gryllidae are so peculiar as the forms we have mentioned.
The family consists in larger part of Insects more or less similar to the
common cricket, though exhibiting a great variety of external form. The
common cricket of our houses, _Gryllus (Acheta) domesticus_ (Fig. 204), has
a very wide distribution in the Old World, and is also found in North
America. It is believed to have had its natural distribution extended by
commerce, though really nothing is known as to its original habitat. The
shrill chirping of this little Insect is frequently heard at night in
houses, even in the most densely inhabited parts of great cities. Neither
the female nor the young are musical, yet the chirping may be heard at all
seasons of the year, as young and adults coexist independent of season. The
predilection of _Gryllus domesticus_ for the habitations of man is very
curious. The Insect is occasionally found out of doors in the neighbourhood
of dwelling-houses in hot weather, but it does not appear that this species
leads anywhere a truly wild life. It is fond of heat; though it rarely
multiplies in dwelling-houses to any great extent, it is sometimes found in
profusion in {339}bake-houses. Usually the wings in the cricket are
elongate, and project backwards from under the tegmina like an additional
pair of cerci; a variety, however, occurs in which these tails are absent,
owing to abbreviation of the wings.
There is no beauty in the appearance of any of the Gryllidae, though many
of them are very bizarre in shape. Very few of them venture to leave the
surface of the earth to climb on plants. The species of _Oecanthus_,
however, do so, and may be found sitting in flowers. They have a more
Locustoid appearance than other Gryllidae. One of the most curious forms of
the family is _Platyblemmus_, a genus of several species found in the
Mediterranean region, the male of which has the head prolonged into a
curious process (Fig. 211); this varies greatly in development in the males
of the same species. It would seem that this organ is of a similar nature
to the extraordinary structures we have figured in Locustidae (Fig. 189)
and Mantidae (Fig. 136), though it appears impossible to treat the cephalic
appendages of _Platyblemmus_ as ornamental objects; their import is at
present quite obscure.
A curious form of variation occurs in this family, and is called
micropterism by de Saussure; we have already mentioned its occurrence in
the house-cricket. The hind wings, which are usually ample, and frequently
have their extremities rolled up and protruding like cerci, are sometimes
much smaller in size, and not visible till the tegmina are expanded. De
Saussure at one time supposed these micropterous individuals to be distinct
species; it is now, however, known that intermediate examples can be found
by examining a great many specimens. Some species are always micropterous.
[Illustration: FIG. 211.—_Platyblemmus lusitanicus_, male. A, front of
head; B, profile of Insect with most of the appendages removed.]
In Britain we have only four representatives of the Gryllidae, viz. the
mole-cricket, the house-cricket, and two field-crickets, one of which,
_Nemobius sylvestris_, is considerably smaller than the house-cricket,
while the other, _Gryllus campestris_, the true field-cricket, is a larger
Insect. Its habits have been described in an interesting manner in Gilbert
White's 88th letter. {340}This Insect, like so many others, is apparently
becoming rare in this country.
A single fossil from the Lias has been described as belonging to the
Gryllidae, but in the Tertiary strata a variety of members of the family
have been discovered both in Europe and North America.
The classification of Gryllidae is due to de Saussure,[267] and is said by
Brunner to be very natural. In the following synopsis of the tribes of
crickets we give de Saussure's arrangement, except that we follow Brunner
in treating Tridactylides as a distinct tribe:—
1. Antennae ten-jointed; posterior tarsi aborted. Tribe 1. TRIDACTYLIDES.
(Fig. 209, _Tridactylus variegatus_; Fig. 210, _Rhipipteryx_ sp.)
1′. Antennae many jointed; posterior tarsi normal.
2. Tarsi compressed, the second joint minute.
3. Anterior legs fossorial; anterior tibiae at the apex with two to
four divisions. Pronotum elongate, ovate, rounded behind. Female
without ovipositor. Tribe 2. GRYLLOTALPIDES. (Fig. 206, front legs of
_Gryllotalpa_; Fig. 208, _Cylindrodes kochi_.)
3′. Anterior legs formed for walking. Ovipositor of the female
visible (either elongate or rudimentary).
4. Posterior tibiae biseriately serrate. Tribe 3. MYRMECOPHILIDES.
4′. Posterior tibiae biseriately spinose. Ovipositor straight.
5. Antennae short, thickish, almost thread-like. Facial
scutellum exserted between antennae. Posterior tibiae dilated.
Gen. _Myrmecophila_.[268]
5′. Antennae elongate, setaceous. Facial scutellum transverse,
visible below the antennae. Tibiae slender.
6. Posterior tibiae armed with two strong spines, not serrate
between the spines. Tribe 4. GRYLLIDES. (Fig. 204, _Gryllus
domesticus_; Fig. 211, _Platyblemmus lusitanicus_.)
6′. Posterior tibiae slender, armed with slender spines, and
serrate between them. Tribe 5. OECANTHIDES.
2′. Second joint of the tarsi depressed, heart-shaped.
3. Posterior tibiae not serrate, but biseriately spinose.
4. The spines on each side three and mobile; apical spurs on the
inner side only two in number. Ovipositor short, curved. Tribe 6.
TRIGONIDIIDES.
4′. The spines numerous, fixed. Ovipositor elongate, straight. Gen.
_Stenogryllus_.
3′. Posterior tibiae serrate and spinose on each side, the apical
spurs, as usual, three on each side. Ovipositor straight or curved.
Tribe 7. ENEOPTERIDES.
{341}CHAPTER XV
NEUROPTERA—MALLOPHAGA—EMBIIDAE
ORDER III. NEUROPTERA.
_Imago with biting mouth; with two pairs of wings, the anterior as well as
the posterior membranous, usually with extensive neuration, consisting of
elongate nervures and either of short cross-nervules forming numerous cells
or of a complex minute mesh-work. (One division, Mallophaga, consists
entirely of wingless forms; in Termitidae some of the individuals of each
generation become winged, but others do not: except in these cases adult
wingless forms are few.) The metamorphosis differs in the several
divisions._
[Illustration: FIG. 212.—_Osmylus chrysops_, New Forest.]
The Neuroptera form a heterogeneous, though comparatively small, Order of
Insects, including termites, stone-flies, dragon-flies, may-flies,
caddis-flies, lace-wings, scorpion-flies, ant-lions, etc. Bird-lice are
also included in Neuroptera, though they have no trace of wings.
We treat the Order as composed of eleven distinct families, {342}and, as a
matter of convenience, arrange them in five divisions:—
1. _Mallophaga_.—Permanently wingless Insects, living on the bodies of
birds or mammals. (Development very imperfectly known.) Fam. 1.
Mallophaga.
2. _Pseudoneuroptera_.—Insects with wings in adult life (in some cases
wings are never acquired). The wings are developed in a visible manner
outside the body. There is no definite pupa. Live entirely on land. Fam.
2. Embiidae; 3. Termitidae; 4. Psocidae.
3. _Neuroptera amphibiotica_.—Wings developed as in division 2. Three
ocelli usually exist. Life aquatic in the early stages. Fam. 5. Perlidae;
6. Odonata; 7. Ephemeridae.
4. _Neuroptera planipennia_.—Wings developed internally; not visible in
early stages, but becoming suddenly evident when the pupal form is
assumed. Mandibles present in the adult Insect. Life in early stages
aquatic or terrestrial. Fam. 8. Sialidae; 9. Panorpidae; 10.
Hemerobiidae.
5. _Trichoptera_.—Development as in division 4. Mandibles absent in the
adult Insect. Life aquatic in the early stages. Fam. 11. Phryganeidae.
The families we have enumerated in the preceding scheme are now generally
adopted by entomologists. Great difference of opinion exists, however, as
to the groups of greater value than the family, and for a long time past
various schemes have been in vogue. Though it is necessary to allude to the
more important of these systems, we can do so only in the briefest manner.
Some of the families of Neuroptera are similar in many points of structure
and development to Insects of other Orders; thus Termitidae are somewhat
allied to Blattidae, Perlidae to Phasmidae in Orthoptera, while the
Phryganeidae or Trichoptera make a considerable approach to Lepidoptera.
Some naturalists—among whom we may mention Burmeister and Grassi—unite our
Aptera, Orthoptera, and most of our Neuroptera into a single Order called
Orthoptera. Others treat our Neuroptera as consisting of eight or nine
distinct Orders; these, together with the names proposed for them, we have
already alluded to in our chapter on classification, pp. 171-177.
Erichson, impressed by the variety existing in Neuroptera, separated some
of the groups into a sub-Order called Pseudoneuroptera; this sub-Order
comprised our Termitidae, Psocidae, Ephemeridae, and Libellulidae. This
division is still adopted in several treatises; the Pseudoneuroptera are
indeed by some naturalists retained as an Order distinct from both
Orthoptera {343}and Neuroptera. Gerstaecker subsequently made use of a
system somewhat different from that of Erichson, uniting the Perlidae,
Ephemeridae, and Odonata into a group called _Orthoptera amphibiotica_,
from which the Termitidae and Psocidae were excluded. The divisions we have
here adopted differ but little from those of Gerstaecker, though we have
arranged them in a very different manner. It is probable that not one-tenth
part of the Neuroptera existing in the world have yet been examined by
entomologists, and of those that are extant in collections, the
life-histories and development are very imperfectly known. We have,
therefore, not considered it wise to adopt a system that would involve
great changes of nomenclature, while there can be little hope of its
permanency.
FOSSILS.—When considering the subject of fossil Insects we briefly alluded
to the discussions that have occurred as to whether the fossils of the
palaeozoic period should be referred to existing Orders. Since the pages we
allude to were printed, M. Brongniart's very important work[269] on the
Insects of that epoch has appeared. He considers that these ancient fossils
may be classified with the existing Orders of Insects, though they cannot
be placed in existing families; and he assigns the palaeozoic fossil
Insects at present known, to the Orders Neuroptera and Orthoptera, and to
the homopterous division of Hemiptera. The greater part of the species he
looks on as Neuroptera, and places in six families—Megasecopterides,
Protephemerides, Platypterides, Stenodictyopterides, Protodonates, and
Protoperlides. Of these he considers the ancient Protephemerides,
Protodonates, and Protoperlides as the precursors, which, we presume, we
may interpret as the actual ancestors, of our existing Ephemeridae,
Odonata, and Perlidae.
Some of the fossils restored and described by the French entomologist are
of great interest. We shall notice the Protephemerides, Protodonates, and
Protoperlides in connexion with the families to which they are specially
allied, and shall now only allude to the quite extinct families of
Neuroptera, the Megasecopterides, Platypterides, and Stenodictyopterides.
It is a peculiarity of these ancient Insects that they were much larger
creatures than the corresponding forms that now exist. This may be due, to
some extent, to the fact that tiny, {344}fragile forms have not been
preserved in the rocks, or have not attracted the attention of collectors;
but as some of the palaeozoic Insects were absolutely the largest
known—surpassing considerably in size any Insects at present existing—it is
probable that, even if small forms existed at the remote epoch we are
alluding to, the average size of the individual was greater than it is at
present. The Megasecopterides of the carboniferous epoch were Insects of
large size, with long, narrow wings, a small prothorax, and large meso- and
meta-thorax, these two segments being equal in size; the abdomen was
elongate and moderately voluminous, and was terminated by a pair of very
elongate, slender filaments like those of the may-flies. The family
includes several genera and species found at Commentry. One of these forms,
_Corydaloides scudderi_, is of great interest, as it is believed by
Brongniart that the imago possessed tracheal gills situated on the sides of
the abdomen, analogous with those that exist at present in the immature
condition of certain Ephemeridae. They are of interest in connexion with
the gills found at the present time in the imagos of _Pteronarcys_ (see p.
401). Although these fossils are of such enormous antiquity, the tracheae
can, M. Brongniart says, be still perceived in these processes.
The Platypterides include also a considerable number of Insects of large
size, with four large equal wings, frequently spotted or variegate. Some of
these Insects were provided with expansions or lobes on the sides of the
prothorax (Fig. 213); these are looked on as analogous to the expansions of
meso- and meta-thorax, which are supposed by some writers to have been the
rudiments from which wings were developed. These prothoracic
wing-rudiments, if such they be, are said to have a system of nervures
similar to what we find in true wings. The genus _Lithomantis_ includes a
Scotch fossil, and has already been mentioned by us on p. 259.
[Illustration: FIG. 213.—_Lithomantis carbonaria_. Carboniferous strata of
Commentry, France. (After Brongniart.)]
The third family of extinct carboniferous Neuroptera is the
Stenodictyopterides, in which Brongniart places the _Dictyoneura_ of
{345}Goldenberg, the North American _Haplophlebium_, and several genera
from Commentry. Some of them were very large Insects, with robust bodies,
and possessed wing-like expansions on the prothorax, and lateral gill-like
appendages on the sides of the abdomen.
It is worthy of note that though so large a number of carboniferous
Neuroptera have now been discovered, no larvae or immature forms have been
found.
We now pass to the consideration of the divisions of Neuroptera still
living.
FAM. I. MALLOPHAGA—BIRD-LICE OR BITING LICE.
_Small Insects, wingless, with large head; thorax usually of two, rarely
of one or three segments; prothorax always distinct; hind body consisting
of eight to ten segments, in addition to the posterior two thoracic
segments which usually are but little or not at all separated from it.
The metamorphosis is very slight. The creatures live on the skins of
birds or mammals, finding nourishment in the epidermal products._
The whole of the Insects of this family live a parasitic, or rather
epizoic, life. They all creep about those parts that are near to the skin,
the feathers of birds or the hair of mammals; they rarely come quite to the
surface, so that they are not detected on a superficial examination. It is
curious that under these circumstances they should exhibit so great a
variety of form and of anatomical characters as they do.
[Illustration: FIG. 214.—_Trinoton luridum_. Lives on the common duck and
various species of _Anas_. (After Giebel.)]
They are very depressed, that is, flat, Insects, with a large head, which
exhibits a great variety of shape; frequently it is provided in front of
the antennae with some peculiar tubercles called trabeculae, which in some
cases are mobile. The antennae are never large, frequently very small; they
consist of from three to five joints, and are sometimes concealed in a
cavity on the under side of the head.
{346}[Illustration: FIG. 215.—Under-surface of head of _Lipeurus
heterographus_. (After Grosse.) _ol_, Labium; _md_, mandible; _mx_,
maxilla; _ul_, labium.]
[Illustration: FIG. 216.—Under lip of _Nirmus_, A; and of _Tetrophthalmus
chilensis_, B. (After Grosse.) _m_, Mentum; _g_, ligula; _pl_, palp; _pg_,
paraglossa; _hy_, lingua.]
The eyes are very rudimentary, and consist of only a small number of
isolated facets placed behind the antennae; sometimes they are completely
absent. The mouth parts are situated entirely on the under-surface of the
head and in a cavity. The upper lip is frequently of remarkable form, as if
it were a scraping instrument (_ol_, Fig. 215). The mandibles are sharply
toothed and apparently act as cutting instruments. The maxillae have been
described in the principal work on the family[270] as possessing in some
cases well-developed palpi. According to Grosse[271] this is erroneous; the
maxillae, he says, are always destitute of palpi, and are of peculiar form,
being each merely a lobe of somewhat conical shape, furnished on one aspect
with hooks or setae. The under lip is peculiar, and apparently of very
different form in the two chief groups of Mallophaga. The large mentum
bears, in Liotheides (Fig. 216, B), on each side a four-jointed palpus, the
pair of palps being very widely separated; the ligula is broad and
undivided; on each side there is a paraglossa bearing an oval process, and
above this is a projection of the hypopharynx. In Philopterides (Fig. 216,
A) the palpi are absent, and the parts of the lower lip are—with the
exception of the paraglossae—but little differentiated. The lingua
(hypo-pharynx) in Mallophaga is largely developed, {347}and bears near the
front a chitinous sclerite corresponding with another placed in the
epipharynx.
The prothorax in Mallophaga is a distinct division of the body even when
the meso- and meta-thorax appear to be part of the abdomen. The mesothorax
is frequently very small; it and the metathorax are sometimes intimately
connected. In other cases (_Laemobothrium_) the metathorax appears to
differ from the following abdominal segment only by having the third pair
of legs attached to it. In _Trinoton_ (Fig. 214) the three thoracic
segments are well developed and distinct. The abdominal segments visible,
vary in number from eight to ten; there is sometimes a difference according
to sex, the male having one segment taken into the interior in connexion
with the reproductive organs. The legs have short, broad coxae and small
tarsi of one or two joints; very rarely three joints are present; there are
either one or two claws; the legs with one claw being adapted for clinging
to or clutching hairs. The front pair of legs is used not for locomotion so
much as for grasping the food and bringing it within the range of the
mouth. No trace of wings has been detected in any species.
[Illustration: FIG. 217.—Ganglia of nervous system of _Lipeurus bacillus_.
(After Giebel.) _a_, Cavity of head.]
The nervous system has been examined by Giebel in _Lipeurus bacillus_;
there is a supra- and an infra-oesophageal ganglion, and three thoracic,
but no abdominal ganglia. The supra-oesophageal is remarkably small, in
fact not larger than the infra-oesophageal; it consists evidently of two
conjoined halves. The alimentary canal has a slender, elongate oesophagus,
dilated behind into a crop; this is frequently received between two cornua
formed by the anterior part of the stomach, which, except for these, is
simply tubular in form, though somewhat narrower at the posterior
extremity. In some forms—Philopterides—the crop is of a very peculiar
nature (Fig. 218), forming an abrupt paunch separated from the stomach by
the {348}posterior portion of the oesophagus. There are only four
Malpighian tubes; in some species the basal half of each tube is much
dilated. The two divisions of the intestine are short and are separated by
the intervention of a glandular girdle. Salivary glands exist; Giebel
figures what we may consider to be an enormous salivary reservoir as
existing in _Menopon leucostomum_.
The testes and ovaries are of a simple nature. The former consist of two or
three capsules, each having a terminal thread; the vasa deferentia are
tortuous and of variable length; they lead into the anterior part of the
ejaculatory duct, where also opens the elongate duct proceeding from the
bicapsular vesicula seminalis; these structures have been figured by
Grosse[272] as well as by Giebel. The ovaries consist of three to five
short egg-tubes on each side; the two oviducts combine to form a short
common duct with which there is connected a receptaculum seminis.
[Illustration: FIG. 218.—Alimentary canal of _Docophorus fuscicollis_.
(After Giebel.) _a_, Oesophagus; _b_, paunch; _a′_, posterior division of
oesophagus; _c_, chylific ventricle or stomach; _d_, Malpighian tubes; _e_,
small intestine; _f_, glandular girdle; _g_, rectum.]
The eggs of some Mallophaga have been figured by Melnikow;[273] they
possess at one extremity a cover with a multiple micropyle-apparatus, and
at the opposite pole are provided with seta-like appendages. They are very
like the eggs of the true lice, and are said in some cases to be suspended
by threads to the hairs or feathers after the fashion of the eggs of
Pediculi.
Little is known as to the development; the young are extremely like the
adult, and are thought to moult frequently; the duration of life is quite
unknown.
It has been stated by some writers that the mouth is truly of the sucking
kind, and that the Mallophaga feed on the blood of their hosts. This is,
however, erroneous; they eat the delicate portions of the feathers of
birds, and of mammals perhaps the young hair. Their fertility is but small,
and it is believed that {349}in a state of nature they are very rarely an
annoyance to their hosts. The majority of the known species live on birds;
the forms that frequent mammals are less varied and have been less studied;
most of them have only one claw to the feet (Fig. 220), while the greater
portion of the avicolous species have two claws.
[Illustration: FIG. 219.—_Lipeurus ternatus_, male; inhabits _Sarcorhamphus
papa_. (After Giebel.)]
[Illustration: FIG. 220.—_Trichodectes latus_, male; inhabits the dog,
_Canis familiaris_.]
Most of the forms have the anterior legs small, and they are usually drawn
towards the mouth, owing, it is believed, to their being used after the
manner of hands to bring the food to the mouth; hence in some of our
figures (219, 220) the body looks as if it had only four legs.
Very diverse statements have been made as to whether allied forms of
Mallophaga are found only on allied birds. It would appear that this is the
case only to a limited extent, as certain species are found on quite a
variety of birds; moreover, some birds harbour several species of
bird-lice, even five genera having been found, it is said, on one species
of bird. _Docophorus icterodes_ has been recorded as occurring on many
kinds of ducks and geese; the swan, however, harbours a distinct species,
_Docophorus cygni_, and this is said to have also been found on the
bean-goose.
At least five species, belonging to three distinct genera, have been found
on the common fowl. The parasite most frequently met with on this valuable
creature is _Menopon pallidum_ (Fig. {350}221), which is said to have been
figured by Redi two hundred years ago under the name of _Pulex capi_. This
species multiplies to a considerable extent; it is of very active habits,
and passes readily from one bird to another, so that it is found on other
species besides the domestic fowl. It is even said that horses kept near
hen-roosts have been seriously troubled by _Menopon pallidum_, but it is
suggested by Osborn that these attacks may perhaps have been really due to
itch-mites. There is, however, no doubt that this species may infest
poultry, especially if sickly, to an enormous extent. The dust-baths in
which poultry are so fond of indulging are considered to be of great use in
keeping down the numbers of this Insect.
[Illustration: FIG. 221.—_Menopon pallidum_; inhabits the common fowl,
_Gallus domesticus_. (After Piaget.)]
A table of the birds and mammals on which Mallophaga have been found,
together with the names of the latter, has been given by Giebel.[274] The
classification of the group, so far as the principal divisions are
concerned, by no means accords with the kind of animals that serve as
hosts, for the only two genera peculiar to quadrupeds (_Trichodectes_, Fig.
220; and _Gyropus_) belong to the two chief divisions of Mallophaga. The
genus _Menopon_ includes numerous species found on birds, and three or four
others peculiar to mammals.
Two very natural divisions, Philopterides and Liotheides, were adopted by
Giebel and Nitzsch, but unfortunately the chief character they made use of
for diagnosing the two groups—the presence or absence of maxillary
palpi—was illusory. Apparently the labial palps will serve the purpose of
distinguishing the two divisions, they being present in the Liotheides and
absent in the Philopterides. A table of the characters of the avicolous
genera of these two groups is given by Grosse.[275]
The Liotheides are more active Insects, and leave their host after its
death to seek another. But the Philopterides do not do so, and die in about
three days after the death of their host. Possibly Mallophaga may be
transferred from one bird to another {351}by means of the parasitic
two-winged flies that infest birds. The writer has recorded[276] a case in
which a specimen of one of these bird-flies captured on the wing was found
to have some Mallophaga attached to it.
We should perhaps point out that these Mallophaga, though called bird-lice,
have nothing to do with the true lice which are so frequently found with
them, and that live by sucking the blood of their hosts. It would in fact
be better to drop the name of bird-lice altogether, and call the Mallophaga
biting lice. _Trichodectes latus_, according to this method, would be known
as the biting louse of the dog, the true or sucking louse of which animal
is _Haematopinus piliferus_, and belongs to the anoplurous division of
Hemiptera.
FAM. II. EMBIIDAE.
_Elongate feeble Insects; with small prothorax, elongate meso- and
meta-thorax, which may either bear wing or be without them. In the former
case these organs are not caducous, are delicately membranous, and all of
one consistence, with three or four indefinite longitudinal nervures and
a few cross-veinlets. The development is incompletely known. The
individuals do not form organised societies._
[Illustration: FIG. 222.—_Oligotoma michaeli._ (After M‘Lachlan.)]
The Embiidae are one of the smallest families of Insects; not more than
twenty species are known from all parts of the world, and it is probable
that only a few hundred actually exist. They are small and feeble Insects
of unattractive appearance, and shrivel so much after death as to render it
difficult to ascertain their characters. They require a warm climate. Hence
{352}it is not a matter for surprise that little should be known about
them.
[Illustration: FIG. 223.—Under-surface of _Embia_ sp. Andalusia.]
The simple antennae are formed of numerous joints, probably varying in
number from about fifteen to twenty-four. The mouth is mandibulate. Chatin
states[277] that the pieces homologous with those of a maxilla can be
detected in the mandible of _Embia_. The labium is divided. The legs are
inserted at the sides of the body, the coxae are widely separated (Fig.
223), the hind pair being, however, more approximate than the others. The
abdomen is simple and cylindrical, consisting of ten segments, the last of
which bears a pair of biarticulate cerci. In the male sex there is a slight
asymmetry of these cerci and of the terminal segment. The thorax is
remarkable on account of the equal development of the meso- and meta-thorax
and their elongation in comparison to the prothorax. When they bear wings
there is no modification or combination of the segments for the purposes of
flight, the condition of these parts being, even then, that of wingless
Insects; so that the Embiidae that have wings may be described as
apterous-like Insects provided with two pairs of inefficient wings. The
wings are inserted on a small space at the front part of each of the
segments to which they are attached. The legs have three-jointed tarsi, and
are destitute of a terminal appendage between the claws.
[Illustration: FIG. 224.—Anterior wing of _Oligotoma saundersii_: A, the
wing; B, outline of the wing, showing nervures. (After Wood-Mason.) 1,
Costal; 2, subcostal; 3, radial; 4, discoidal; 5, anal nervure.]
The wings in Embiidae are very peculiar; they are extremely {353}flimsy,
and the nervures are ill-developed; stripes of a darker brownish colour
alternate with pallid spaces. We figure the anterior wing of _Oligotoma
saundersii_, after Wood-Mason; but should remark that the neuration is
really less definite than is shown in these figures; the lower one
represents Wood-Mason's interpretation of the nervures. He considers[278]
that the brown bands "mark the original courses of veins which have long
since disappeared." A similar view is taken by Redtenbacher,[279] but at
present it rests on no positive evidence.
One of the most curious features of the external structure is the complex
condition of the thoracic sternal sclerites. These are shown in Fig. 223,
representing the under-surface of an _Embia_ of uncertain species recently
brought by Mr. Bateson from Andalusia.
According to Grassi[280] there are ten pairs of stigmata, two thoracic and
eight abdominal; these are connected by longitudinal and transverse
tracheae into a single system. The ganglia of the ventral chain are, one
suboesophageal, three thoracic, and seven abdominal; these are segmentally
placed, except that there is no ganglion in the fifth abdominal segment.
There is a stomato-gastric system but no "sympathetic." Salivary glands are
present. The stomodaeal portions of the alimentary canal are remarkably
capacious; the stomach is elongate and slender, without diverticula; the
Malpighian tubes are elongate and slender; they vary in number with the age
of the individual, attaining that of twenty in the adult. The ovaries are
arranged somewhat after the fashion of those of _Japyx_, there being in
each five short egg-tubes, opening at equal intervals into a straight duct.
The testes are remarkably large; each one consists of five masses of
lobules, and has a large vesicula seminalis, into the posterior part of
which there open the ducts of two accessory glands. The large joint of the
front tarsus includes glands whose secretion escapes by orifices at the
tips of certain setae interspersed between the short spines that are placed
on the sole.
Species of this genus occur in the Mediterranean region, but their
characters have not yet been examined. Our information {354}as to these is
chiefly to be found in Grassi's work. The two species studied by him were
wingless. They live under stones, where they spin webs by means of the
front feet, whose first joint is, as we have said, enlarged and contains
glands; the Insect uses the webs as a means of support in progression,
acting on them by means of papillae and a comb-like structure placed on the
four posterior feet.
Grassi informs us that these Insects are not uncommon under stones in
Catania; they require moisture as well as warmth, but not too much;
sometimes there is only one individual found under a stone, at others eight
or ten. In the winter and spring their galleries are found on the surface
of the earth, but in the hot months of summer they secure the requisite
amount of moisture by sinking their galleries to the depth of ten or
fifteen centimetres. Their food consists chiefly of vegetable matter. They
may be reared with ease in glass vessels. Other species of the family
attain wings; the details of the process are not well known. _Oligotoma
michaeli_ (Fig. 222) was discovered in a hothouse in London among some
orchid roots brought from India, and was found in more than one stage of
development; the young greatly resemble the adult, except in the absence of
wings. A nymph-form is described by M‘Lachlan[281] as possessing wings of
intermediate length, and Hagen has suggested that this supposed nymph is
really an adult female with short wings. If this latter view be correct,
nothing is known as to the mode of development of wings in the family. It
is still uncertain whether female Embiidae ever possess wings. Wood-Mason
and Grassi have shown that there are wingless females in some species, and
we know that there are winged males in others, but what the usual relation
of the sexes may be in this respect is quite uncertain. These Insects have
been detected in various parts of the world. In the Sandwich Islands
_Oligotoma insularis_ was discovered by the Rev. T. Blackburn in the wood
and thatch forming the roofs of natives' houses. A species has been found
in Prussian amber, and Grassi thinks that _Embia solieri_—one of the
Mediterranean species—is not to be distinguished with certainty from the
Insect found in amber.
Embiidae still remains one of the most enigmatic of the families of
Insects. Although Grassi's recent observations are {355}of great value from
an anatomical point of view, they rather add to, than diminish, the
difficulties we encounter in endeavouring to understand the lives of these
obscure creatures. That Embiidae form webs has long been known, and it was
thought by some that the webs, like those of spiders, might be of
assistance in procuring food. We may, however, infer from Grassi's
observations that this is not the case, but that the silken tunnels or
galleries—as he calls them—serve chiefly as a means of locomotion and
protection, the feet of the Insects being highly modified in conformity
with this mode of life. Grassi seems to be of opinion that the galleries
are also useful in preserving a proper degree of humidity round the
Insects. We have already alluded to the mystery that surrounds the mode of
growth of their wings. Nearly all that is known as to the Embiidae is
contained in Grassi's paper, or is referred to in Hagen's monograph of the
family.[282]
Considerable difference of opinion has prevailed as to the allies of these
obscure Insects. It would seem that they are most nearly allied to
Termitidae and Psocidae. Grassi, however, considers these affinities only
remote, and suggests that Embiidae should form a separate Order, to be
placed in a super-Order Orthoptera, which would include our Aptera, the two
families mentioned above, Mallophaga, Embiidae, and the ordinary
Orthoptera. Brauer places the family in his Orthoptera genuina.
{356}CHAPTER XVI
NEUROPTERA _CONTINUED_—TERMITIDAE, TERMITES OR WHITE ANTS
FAM. III. TERMITIDAE—WHITE ANTS, TERMITES.
[Illustration: FIG. 225.—_Termes_ (_Hodotermes_) _mossambicus_. Winged
adult. (After Hagen.)]
_Each species is social, and consists of winged and wingless individuals.
The four wings are, in repose, laid flat on the back, so that the upper
one only is seen except just at the bases; they are membranous and very
elongate, so that they extend far beyond the apex of the abdomen; the
hind pair is remarkably similar in size, form, and consistence to the
front pair: near the base of each wing there is a suture, or line of
weakness, along which the wings can be broken off, the stumps in that
case remaining as short horny flaps reposing on the back. Ligula
channelled but not divided into two parts. The wingless individuals are
very numerous, and have the head and thirteen body segments distinct; the
body {357}is terminated by a pair of short cerci. The metamorphosis is
slight and gradual, and in some individuals is dispensed with._
The term White Ants has been so long in use for the Termitidae that it
appears almost hopeless to replace it in popular use by another word. It
has, however, always given rise to a great deal of confusion by leading
people to suppose that white ants differ chiefly from ordinary ants by
their colour. This is a most erroneous idea. There are scarcely any two
divisions of Insects more different than the white ants and the ordinary
ants. The two groups have little in common except that both have a social
life, and that a very interesting analogy exists between the forms of the
workers and soldiers of these two dissimilar Orders of Insects, giving rise
to numerous analogies of habits. The word Termites—pronounced as two
syllables—is a less objectionable name for these Insects than white ants.
The integument in Termites is delicate, and the chitinous plates are never
very hard; frequently they are so slightly developed that the creature
appears to consist of a single membranous sac with creases in it, the head
alone being very distinct. The head is exserted, frequently of large size,
sometimes as large as all the rest of the body together. Termites may be
quite blind, or possess facetted and simple eyes, the latter when present
being two in number and always accompanied by facetted eyes. The antennae
are simple, consisting of from nine to thirty-one joints, which differ but
little from one another; the number in each individual increases as the
development progresses. The parts of the mouth are large, the ligula
consists of one piece (Fig. 226, A), but often has the appearance of being
formed by two united pieces; on its extremity are seated two pairs of
lobes.
[Illustration: FIG. 226.—_Termes bellicosus._ Labium, A, maxilla, B, of
winged adult; lower face of each. (After Hagen.)]
The head is articulated to the thorax by means of two very large cervical
sclerites on each side, placed at right angles to one another, and visible
on the under-surface. The prothorax is well developed and distinct from the
parts behind it. The {358}pronotum, of variable form and size, is very
distinct in the perfect Insects; with it are connected the largely
developed pleura. The episternum is very peculiar, consisting of an
elongate chitinous slip on each side hanging downwards, the two not quite
meeting in the middle; they thus form the margin of the very large anterior
orifice, and are in contiguity with the cervical sclerites; behind them are
the very large epimera. The prosternum appears to be usually entirely
membranous; in some cases the sclerite in it is small and delicate, and
apparently differs according to the species. The meso- and meta-thorax are
sub-equal in size; the mesosternum forms a peculiar, large, adpressed fold.
The metasternum is membranous, but is terminated behind by a sclerite
apparently of variable form. The hind body is voluminous, simple in form,
consisting of ten segments and bearing at the extremity two short distant
cerci of a variable number of joints. The terminal ventral sclerites differ
greatly in form according to the species and sometimes according to the
sex; there are sometimes, if not always, present near the extremity two
peculiar minute biarticulate styles, called appendices anales. The coxae
are all large, free, and exserted; at the base of each is a transverse
trochantin. The femora are articulated with the trochanters, not with the
coxae; both femora and tibiae are slender, the tarsi small, four-jointed;
the terminal joint elongate.
[Illustration: FIG. 227.—Front tibia and tarsus of _Calotermes rugosus_
larva, showing auditory organ. × 90. (After F. Müller.)]
It is now well established that Termites have a means of communication by
sounds. The individuals have a peculiar way of jerking themselves, as has
been frequently noticed by observers of the Insects; these convulsive
movements may possibly be connected with the production of sound, which may
perhaps be evoked by contact between the back of the head and the pronotum;
the exact mode by which the sounds are produced is not, however, known. The
existence of an auditory organ in the front tibia has been demonstrated by
Fritz Müller,[283] and we reproduce (Fig. 227) one of his figures. The
structure seems to {359}be in plan and position similar to the ear of
Locustidae, though much less perfect.
[Illustration: FIG. 228.—Wings of Termites: A, _Termes lucifugus_; B,
_Hodotermes brunneicornis_; C, _Culotermes nodulosus_. (After Redtenbacher:
B and C diagrammatic.) III, V, VII, homologous areas and nervures according
to Redtenbacher. 1, Costal; 2, subcostal; 3, median; 4, submedian nervures
according to Hagen.]
The wings of Termitidae are not like those of any other Insects; their
neuration is very simple, but nevertheless the wings of the different forms
exhibit great differences in the extent to which they are made up of the
various fields. This is shown in Fig. 228, where the homologous nervures
are numbered according to the systems of both Hagen and Redtenbacher. The
area, VII, that forms the larger part of the wing in C, corresponds to the
small portion at the base of the wing in B. The most remarkable feature of
the wing is, however, its division into two parts by a suture or line of
weakness near the base, as shown in Fig. 225. The wings are used only for a
single flight, and are then shed by detachment at this suture; the small
basal portion of each of the four wings is horny and remains attached to
the Insect, serving as a protection to the dorsal surface of the thorax.
The nature of the suture that enables the Termites to cast their wings with
such ease after swarming is not yet understood. There are no true
transverse veinlets or nervules in Termites. Redtenbacher suggests[284]
that the transverse division of the wing at its base, as shown in Fig. 225,
along which the separation of the wing occurs at its falling off, may have
arisen from a coalescence of the subcostal vein with the eighth concave
vein of such a wing as that of Blattidae. The same authority also informs
us that the only point of resemblance between the wings of Termitidae and
those of Psocidae is that both have an unusually small number of concave
veins.
The information that exists as to the internal anatomy of {360}Termites is
imperfect, and refers, moreover, to different species; it would appear that
considerable diversity exists in many respects, but on this point it would
be premature to generalise. What we know as to the respiratory system is
chiefly due to F. Müller.[285] The number of spiracles is ten; Hagen says
three thoracic and seven abdominal, Müller two thoracic and eight
abdominal. In fertile queens there usually exist only six abdominal
stigmata. There is good reason for supposing that the respiratory system
undergoes much change correlative with the development of the individual;
it has been suggested that the supply of tracheae to the sexual organs is
deficient where there is arrest of development of the latter.
The alimentary canal is only of moderate length. Salivary glands exist, as
also do salivary reservoirs; these latter are large, in some species
remarkably so. The oesophagus is slender, but abruptly enlarged behind to
form a large crop; a proventriculus is apparently either present or absent;
the chylific ventricle, or stomach, is slender and simple. The Malpighian
tubules are very long; their number is probably from four to eight in the
adult, and in the earlier stages less. Behind the tubes the alimentary
canal forms a large paunch, and after this there is a small intestine and
rectum. The paunch is a peculiar structure, and probably of great
importance in the economy of Termites.
These creatures emit minute quantities of a secretion that is corrosive,
and can act on metal and even glass;[286] its nature and source are not
understood. Hagen describes peculiar structures in the rectum to which he
is inclined[287] to ascribe the origin of this substance, but this is very
uncertain.
[Illustration: FIG. 229.—Head and alimentary canal of _Termes lucifugus_
(nymph). _a_, head; _b_, salivary glands; _c_, salivary receptacles; _d_,
crop; _e_, stomach; _f_, intestinal paunch; _g_, small, _h_, large
intestine; _i_, Malpighian tubes; _k_, extremity of body. (After Dufour.)]
The brain is small; the infra-oesophageal ganglion is placed
{361}immediately under the supra-oesophageal; there are three thoracic and
six abdominal ganglia. The nervous system apparently differs but little in
the various forms, or in the different stages of life, except that in the
fertile females the abdominal ganglia become so much enlarged that they
even exceed the brain in size.
The testes are unusually simple; each consists of eight capsules opening
into the vas deferens; the two vasa converge and are continued as a short
ejaculatory duct; at the point of convergence there is a pair of curled
vesiculae seminales.
The ovarian system is also simple; there is a variable number of elongate
egg-tubes, each of which opens separately into the oviduct; the two ducts
unite to form a short uterus, on which there is placed first a spermatheca,
and near the extremity a convolute tubular sebific gland. The number of
egg-tubes is subject to extraordinary variation, according to the species,
and according to the age of the fertilised individual.
SOCIAL LIFE.—Termites live in communities that consist sometimes of
enormous numbers of individuals. The adult forms found in a community are
(1) workers; (2) soldiers; (3) winged males and females; (4) some of these
winged forms that have lost their wings. Some species have no worker caste.
The individuals of the third category are only present for a few days and
then leave the nest in swarms. In addition to the adult individuals there
are also present various forms of young. The individuals that have lost
their wings are usually limited to a single pair, king and queen; there may
be more than one king and queen, but this is not usual. The king and queen
may be recognised by the stumps of their cast wings, which exist in the
form of small triangular pieces folded on the back of the thorax (Fig.
235). The continuance of the community is effected entirely by the royal
pair; they are the centres of activity of the community, which is thrown
into disorder when anything happens to them. Usually the pair are
physically incapable of leaving the nest, especially the queen, and
frequently they are enclosed in a cell which they cannot leave. In
consequence of the disorganisation that arises in the community in the
absence of a royal pair, Termites keep certain individuals in such a state
of advancement that they can rapidly be developed into royalties should
occasion require it. These reserve individuals are called complementary by
Grassi; when {362}they become royalties they are usually immature as
regards the condition of the anterior parts of the body, and are then
called by Grassi and others neoteinic, as is more fully explained on p.
380.
SWARMS.—As a result of the Termite economy large numbers of superfluous
individuals are frequently produced; these, in the winged state, leave the
community, forming swarms which are sometimes of enormous extent, and are
eagerly preyed on by a variety of animals including even man. Hagen has
given particulars[288] of a swarm of _T. flavipes_ in Massachusetts, where
the Insects formed a dark cloud; they were accompanied by no less than
fifteen species of birds, some of which so gorged themselves that they
could not close their beaks.
There is but little metamorphosis in Termitidae. Young Termites are very
soft; they have a thin skin, a disproportionately large head, and are of a
peculiar white colour as if filled with milk. This condition of milkiness
they retain, notwithstanding the changes of form that may occur during
their growth, until they are adult. The wings first appear in the form of
prolongations of the meso- and meta-nota, which increase in size, the
increment probably taking place at the moults. The number of joints of the
antennae increases during the development; it is effected by growth of the
third joint and subsequent division thereof; hence the joints immediately
beyond the second are younger than the others, and are usually shorter and
altogether more imperfect. The life-histories of Termites have been by no
means completely followed; a fact we can well understand when we recollect
that these creatures live in communities concealed from observation, and
that an isolated individual cannot thrive; besides this the growth is, for
Insects, unusually slow.
NATURAL HISTORY.—The progress of knowledge as to Termites has shown that
profound differences exist in the economy of different species, so that no
fair general idea of their lives can be gathered from one species. We will
therefore briefly sketch the economy, so far as it has been ascertained, in
three species, viz. _Calotermes flavicollis_, _Termes lucifugus_, and _T.
bellicosus_.
{363}[Illustration: FIG. 230.—Some individuals of _Calotermes flavicollis_:
A, nymph with partially grown wing-pads; B, adult soldier; C, adult winged
individual. (After Grassi.)]